Gymnaconitum, a New Genus of Ranunculaceae Endemic to the Qinghai-Tibetan Plateau

Home , Aconitum anthora, Aconitum carmichaelii, Aconitum columbianum, Aconitum ferox, Aconitum flavum, Aconitum heterophyllum

TAXON 62 (4) • August 2013: 713–722 Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

Gymnaconitum, a new genus of Ranunculaceae endemic to the Qinghai-Tibetan Plateau

Wei Wang,1 Yang Liu,2 Sheng-Xiang Yu,1 Tian-Gang Gao1 & Zhi-Duan Chen1

1 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, P.R. China 2 Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut 06269-3043, U.S.A. Author for correspondence: Wei Wang, [email protected]

Abstract The monophyly of traditional Aconitum remains unresolved, owing to the controversial systematic position and taxonomic treatment of the monotypic, Qinghai-Tibetan Plateau endemic A. subg. Gymnaconitum. In this study, we analyzed two datasets using maximum likelihood and Bayesian inference methods: (1) two markers (ITS, trnL-F) of 285 Delphinieae species, and (2) six markers (ITS, trnL-F, trnH-psbA, trnK-matK, trnS-trnG, rbcL) of 32 Delphinieae species. All our analyses show that traditional Aconitum is not monophyletic and that subgenus Gymnaconitum and a broadly defined Delphinium form a clade. The SOWH tests also reject the inclusion of subgenus Gymnaconitum in traditional Aconitum. Subgenus Gymnaconitum markedly differs from other species of Aconitum and other genera of tribe Delphinieae in many non-molecular characters. By integrating lines of evidence from molecular phylogeny, divergence times, morphology, and karyology, we raise the monotypic A. subg. Gymnaconitum to generic status.

Keywords Aconitum; Delphinieae; Gymnaconitum; monophyly; phylogeny; Qinghai-Tibetan Plateau; Ranunculaceae; SOWH test

Supplementary Material The Electronic Supplement (Figs. S1–S8; Appendices S1, S2) and the alignment files are available in the Supplementary Data section of the online version of this article (http://www.ingentaconnect.com/content/iapt/tax).

Received: 30 Sep. 2012; revision received: 22 Apr. 2013; accepted: 30 June 2013. DOI: http://dx.doi.org/10.12705/624.10

INTRODUCTION (DC.) Peterm. is the earliest-diverging lineage within the tribe (Jabbour & Renner, 2011a, 2012a) and has been raised Ranunculaceae represent one of the earliest-diverging to generic status (Jabbour & Renner, 2011b). Thus, Jabbour lineages among the eudicots (APG III, 2009; Sun & al., 2011) & Renner (2012a) considered that tribe Delphinieae contained and consists of 59 genera with about 2500 species (Tamura, three genera: Aconitum, Delphinium (containing Aconitella 1995). The family contains many genera of pharmaceutical and Consolida), and Staphisagria J. Hill. However, the mono- and horticultural interest, such as Aquilegia L., Coptis Salisb. phyly of Aconitum remains unresolved. This genus comprises and Delphinium L. (W.T. Wang, 1979; Peng & al., 2006). In the about 300 species distributed in three subgenera, A. subg. past two decades, tremendous progress has been made in our Lycoctonum (DC.) Peterm., subg. Aconitum (Stapf.) Rapa- understanding of phylogenetic relationships in Ranunculaceae ics, and subg. Gymnaconitum (Stapf.) Rapaics (Tamura, at various taxonomic levels by using DNA sequence data. The 1995). Aconitum subg. Gymnaconitum contains only one subfamily- and tribe-level classification system of Ranuncul species, A. gymnandrum Maxim. (Fig. 1), and is endemic aceae has recently been updated (W. Wang & al., 2009), with to the Qinghai-Tibetan Plateau (Fig. 2). Based on root char- recognition of five subfamilies: Glaucidioideae, Hydrastid acters and life cycle, Stapf (1905) first erected a section for oideae, Coptidoideae, Thalictroideae, and Ranunculoideae this species. Rapaics (1907) further raised it to subgeneric (including ten tribes). Meanwhile, twelve genera accepted by rank, which has been accepted by all subsequent authors (e.g., Tamura (1995) have been synonymized with six other genera, W.T. Wang, 1979; Tamura, 1995). Molecular phylogenetic and four genera not recognized by Tamura (1995) have been studies at the population level (L. Wang & al., 2009) or sam- reinstated based on molecular data (Table 1). The circumscrip- pling multiple accessions of the species (Jabbour & Renner, tions of only a few genera in Ranunculaceae remain contro- 2012a) have indicated that A. gymnandrum is monophyletic. versial and need to be further clarified, such as Aconitum L. The species exhibits some unique traits, such as clawed sepals Aconitum is a member of tribe Delphinieae, which also (Fig. 1B), large fan-shaped petaline labia (Fig. 1C), exposed contains Aconitella Spach, Consolida (DC.) S.F. Gray, and stigmas (Fig. 1B, C), and many carpels (Fig. 1D), which are Delphinium. Based on molecular evidence, Aconitella and not found in other species of Aconitum and other genera of Consolida are embedded within Delphinium (Luo, 2003; Jab- tribe Delphinieae (W.T. Wang, 1979; Tamura, 1995). Thus, bour & Renner, 2011a, 2012a); Delphinium subg. Staphisagria the taxonomic position of the subgenus is a key element in

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determining the monophyly of Aconitum (Xiao & al., 1984, MATERIALS AND METHODS 2006). The phylogenetic position of A. subg. Gymnaconitum has been investigated with various non-molecular charac- Data matrices. — Two data matrices were constructed ters, including phytochemical (Xiao & al., 1984), cytological for phylogenetic analyses. (1) A two-marker dataset (ITS and (Shang, 1985), anatomical (Li & al., 1991; Yang, 1990), as well trnL-F) which is an extension of the recent two-marker dataset as palynological characters (Xi, 1993). However, the subgenus of Jabbour & Renner (2012a), which included 185 species of has both primitive and advanced features in terms of phyto- tribe Delphinieae. Our present matrix sampled 285 species of chemistry and morphology (Xiao & al., 2006). Luo (2003) the tribe, of which 97 are of ca. 300 of Aconitum, 161 are of ca. first used molecular data (plastid trnL-F and nuclear ITS) to 320 of Delphinium, 6 are of ca. 10 of Aconitella, and 21 are of investigate the monophyly of Aconitum, where 23 species of ca. 40 of Consolida (Tamura, 1995; Electr. Suppl.: Appendix S1). tribe Delphinieae (Aconitum represented by 16 species) were For Aconitum and Delphinium, we followed the subgeneric clas- sampled and Delphinium subg. Staphisagria was not included. sifications of Tamura (1995). In Aconitum, subgenus Lycoctonum In her neighbour joining analysis, Aconitum subg. Gymnaco- contains ca. 40 species, of which 18 were included, subgenus nitum was sister to the clade including A. subg. Lycoctonum Aconitum contains ca. 250 species, of which 78 were included, and subg. Aconitum, Delphinium and Consolida with weak and A. gymnandrum as the only species of subgenus Gymnaconi- support (Luo, 2003). Based on plastid (trnL-F, trnK-matK, tum was included. Delphinium consists of three subgenera, subg. trnS-trnG) and nuclear (ITS) sequences, Jabbour & Renner Delphinastrum (DC.) Peterm. (ca. 300 spp.), subg. Staphisagria (2011a) found a monophyletic Aconitum (represented by eight (DC.) Peterm. (3 spp.), and subg. Delphinium W.T. Wang (ca. species) with 55% bootstrap support and resolved A. subg. 18 spp.), of which 143, 3, and 15 species were included, respec- Gymnaconitum as basalmost in Aconitum. However, A. subg. tively. (2) The second dataset was a six-marker dataset (ITS, Gymnaconitum was identified as the second-diverging lineage trnL-F, trnH-psbA, trnK-matK, trnS-trnG, rbcL) containing 32 in tribe Delphinieae with poor support by using trnL-F and Delphinieae species with at least four of the above sequences ITS data and sampling 185 species of the tribe, of which 57 (Electr. Suppl.: Appendix S2). Although taxon sampling density were from Aconitum (Jabbour & Renner, 2012a). Importantly, in this dataset was lower than that in the two-marker dataset, all when the taxon sampling was reduced from 185 to 73 spe- representative genera and subgenera of Tamura (1995) and major cies of tribe Delphinieae, A. subg. Gymnaconitum was sup- lineages of Jabbour & Renner (2012a) in tribe Delphinieae were ported as sister to a broadly defined Delphinium (including included. Following W. Wang & al. (2009), we selected Nigella Consolida and Aconitella; Jabbour & Renner, 2012a). Thus, damascena L. (Nigelleae) and Megaleranthis saniculifolia Ohwi these studies indicate that different taxon and character sam- (Adonideae) as outgroups for these two datasets. The study did pling schemes can affect the phylogenetic position of A. subg. not generate new data, and all DNA sequences were obtained Gymnaconitum. from GenBank. The authorities, herbarium vouchers, localities, In this study, two different data matrices were constructed and accession numbers of species sampling are listed in Appen- with more extensive taxon and character sampling schemes. dices S1 and S2 (Electr. Suppl.). Using these data, our goals are to investigate the monophyly Phylogenetic analysis. — Sequences were aligned using of Aconitum, and then to clarify the phylogenetic position and the default parameters in Clustal X v.1.83 (Thompson & al., taxonomic treatment of A. subg. Gymnaconitum in tribe Del- 1997) and manually adjusted with BioEdit v.5.0.9 (Hall, 1999). phinieae. For the six-marker dataset, phylogenetic analyses were initially

Table 1. Newly delimited or resurrected genera after Tamura (1995). Newly delimited or recognized genera Genera of Tamura (1995) Studies Actaea Actaea, Souliea, Cimicifuga Compton & al. (1998) Anemone Anemone, Hepatica, Pulsatilla, Knowltonia, Oreithales, Barneoudia Hoot & al. (2012, and references therein) Clematis Clematis, Archiclematis, Naravelia Xie & al. (2011) Coptidium – Emadzade & al. (2010) Delphinium Delphinium, Consolida Jabbour & Renner (2011a, 2012a) Ficaria – Emadzade & al. (2010) Paropyrum – W. Wang & Chen (2007) Ranunculus Ranunculus, Aphanostemma Emadzade & al. (2010) Staphisagria – Jabbour & Renner (2011b) Trollius Trollius, Megaleranthis W. Wang & al. (2010)

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Fig. 1. Aconitum gymnandrum. A, plants and habitat; B, frontal view of a flower showing upper sepal (a), lateral sepals (b), and lower sepals (c); C, lateral view of a flower showing petals (d); D, carpels; E, seeds. — Scale bars: A = 10 cm; B, C = 1 cm; D = 1.5 cm; E = 5 mm.

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Fig. 2. Geographical distribution of Aconitum gymnandrum (solid circles).

conducted for individual regions of the cpDNA data using replicate datasets using Seq-Gen v.1.3.2 (Rambaut & Grassly, maximum likelihood (ML). No significant bootstrap support 1997). For each simulated dataset, ML searches were conducted (exceeding 70%) for conflicting nodes was evident among in- fully optimized or under constrained conditions. The lnL dif- dividual chloroplast markers, five plastid datasets were thus ferences (ΔlnL) were then calculated and used for evaluating combined (referred to as cpDNA data). Detailed analyses were the significance of the difference between the best tree and the conducted using ML and Bayesian inference (BI) for the com- constrained tree based on the original data. A detailed descrip- bined two- and six-marker datasets as well as for each of the tion of the SOWH test can be found in Liu & al. (2012). two genome datasets. The ML and BI analyses were conducted in RAxML v.7.2.6 (Stamatakis, 2006) and MrBayes v.3.1.2 (Ronquist & Huelsenbeck, 2003), respectively. RESULTS RAxML was conducted with the GTR + Γ substitution model for each region, and the fast bootstrap option, using 1000 Two-marker dataset. — The aligned trnL-F sequences replicates. For BI analyses, each DNA region was assigned its were 1512 nucleotides long. We excluded 42 ambiguous sites own model of nucleotide substitution, as determined by the located in one region (804–845) from the phylogenetic analy- Akaike information criterion (AIC) in Modeltest v.3.7 (Posada ses. The tree generated by the ML analysis (Electr. Suppl.: & Crandall, 1998). Four Markov chain Monte Carlo chains were Fig. S1) was highly congruent with those retrieved with the BI run, sampling one tree every 1000 generations for 50,000,000 analysis (Electr. Suppl.: Fig. S2), except for containing some generations, starting with a random tree. Stationarity of the weakly supported nodes (BS < 70%). The aligned matrix of ITS runs was assessed using Tracer v.1.5 (Rambaut & Drummond, sequences had a length of 681 characters. ML and BI analyses 2009). A majority-rule (> 50%) consensus tree was constructed resulted in highly congruent trees (Electr. Suppl.: Figs. S3, S4). after removing the burn-in period samples (the first 25% of Relationships among the genera and subgenera revealed from sampled trees). Posterior probabilities (PP) were estimated by the trnL-F data were consistent with those from the ITS data. sampling trees from the PP distribution. The combined trnL-F and ITS dataset consisted of 2151 char- Swofford-Olsen-Waddell-Hillis test. — We used a para- acters, obtained from 287 accessions. The ML tree is shown in metric bootstrapping likelihood ratio test, the Swofford-Olsen- Fig. 3 and in Fig. S5 (Electr. Suppl.). Delphinium subg. Staph- Waddell-Hillis (SOWH) test (Goldman & al., 2000), to assess isagria is the earliest-diverging lineage in tribe Delphinieae. the monophyly of traditional Aconitum. The SOWH test has Aconitum subg. Aconitum and subg. Lycoctonum form a clade been found to have more power and lower occurrence of type I (here named Aconitum s.str.) with strong support (BS 99%, PP error than other hypothesis testing methods when model param- 1.00). Aconitum subg. Gymnaconitum is sister to the clade con- eters are accurately provided (Goldman & al., 2000; Buckley, taining Consolida (sensu Tamura, 1995; including Aconitella) 2002). For each of the two combined datasets, we first con- and Delphinium subg. Delphinium and subg. Delphinastrum strained all taxa of traditional Aconitum as monophyletic, and (BS 65%, PP 0.97). The monophyly of D. subg. Delphinium is then optimized the tree in RAxML (GTR + Γ model, partitioned not supported. Relationships among the genera and subgenera by DNA region). Based on tree topology, branch lengths and revealed by the BI analyses (Electr. Suppl.: Fig. S6) were identi- tree scores generated from the above step, we simulated 100 cal to those from the ML analysis.

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Six-marker dataset. — The aligned trnL-F, trnH-psbA, significant at the 0.05 level (9.0 units). The six-marker dataset trnK-matK, and trnS-trnG matrices comprised 1331, 436, yielded a difference of 8.9 units, which is highly significant 1548, and 960 nucleotides, respectively. Correspondingly, one at the 0.01 level (4.0 units; Fig. 5). The SOWH test indicated difficult-to-align region encompassing 38 sites (696–733) in that the monophyly of traditional Aconitum should be rejected. the trnL-F dataset, one encompassing 50 sites (387–436) in the trnH-psbA dataset, three encompassing 56 sites (277–291, 432–440, and 680–711) in the trnK-matK dataset, and two en- DISCUSSION compassing 45 sites (532–540 and 652–687) in the trnS-trnG dataset were excluded from the analyses. The aligned ITS and All analyses based on our two- and six-marker datasets rbcL datasets had 670 and 1262 positions, respectively. Rela- support the division of tribe Delphinieae into four major tionships within tribe Delphinieae revealed by the cpDNA data clades, Staphisagria, Aconitum s.str., A. subg. Gymnaconi- (Electr. Suppl.: Fig. S7) were identical with those from the ITS tum, and a broadly defined Delphinium (including Consolida data (Electr. Suppl.: Fig. S8). The combined six-marker dataset and Aconitella), with Staphisagria as sister to the remaining consisted of 6018 characters, obtained from 34 accessions. ML Delphinieae. Subgenera Aconitum and Lycoctonum of Aconi- and BI analyses resulted in identical topologies (Fig. 4). Pat- tum s.str., Delphinium subg. Delphinastrum, and Consolida terns of relationships at the generic and subgeneric levels were (sensu Tamura, 1995; including Aconitella) are all strongly identical with those obtained from the two-marker dataset, but supported as monophyletic. These results are consistent with support for Aconitum subg. Gymnaconitum as sister to the clade previous studies (Jabbour & Renner, 2011a, 2012a). Whereas containing Consolida and Delphinium subg. Delphinium and Jabbour & Renner (2011a) found D. subg. Delphinium to be subg. Delphinastrum increased notably (BS 73%, PP 1.00). monophyletic with poor bootstrap support, our two- and six- SOWH test. — For the two-marker dataset, the SOWH marker datasets indicate that Consilida (sensu Tamura, 1995) test indicated that constraining traditional Aconitum as mono- is embedded within D. subg. Delphinium with weak support phyletic yielded an ln-likelihood that is 13.0 units worse than (Figs. 3, 4), which is congruent with the result of Jabbour the unconstrained optimal tree (Fig. 5). This difference is & Renner (2012a). Importantly, our analyses further indicate

Tamura Jabbour & Renner (1995) (2012) 100 Consolida (27/ca. 50) 1.00

99 1.00

99 Delphinium subg. Delphinastrum (143/ca. 300) Delphinium 1.00 49 0.69

99 1.00 65 100 0.97 1.00 Delphinium subg. Delphinium (15/ca. 17)

Aconitum subg. Gymnaconitum (1/1) 96 1.00 98 1.00 Aconitum subg. Aconitum (78/ca. 250) Aconitum 100 99 1.00 1.00 100 1.00 Aconitum subg. Lycoctonum (18/ca. 40) 100 1.00 Delphinium subg. Staphisagria (3/3) Staphisagria Nigella damascena Megaleranthis saniculifolia Outgroups 0.02

Fig. 3. Phylogenetic relationships in tribe Delphinieae obtained from an ML analysis of the two-marker dataset. Numbers above and below branches are bootstrap values and Bayesian posterior probabilities, respectively. Numbers alongside generic or subgeneric names represent the proportion of species we sampled relative to the total species number of each group. Classifications by Tamura (1995) and Jabbour & Renner (2012a) are shown on the right. For the expanded version see Fig. S5 in the Electronic Supplement.

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that traditional Aconitum is not monophyletic, and that A. subg. lineages, i.e., Staphisagria, A. gymnandrum, Aconitum s.str., Gymnaconitum and Delphinium (sensu Jabbour & Renner, and Delphinium, in a narrow window of time (32.3–27.24 Ma). 2012a) form a clade (Figs. 3, 4). To further clarify the phyloge- Thirteen accessions (ITS) of A. gymnandrum grouped together netic position and taxonomic status of A. subg. Gymnaconitum and supported this species as monophyletic (Jabbour & Renner, in tribe Delphinieae, we present molecular, morphological and 2012a). Sampling 23 populations, L. Wang & al. (2009) found karyological evidence. that A. gymnandrum only began to diversify at the population Molecular evidence. — Our phylogenetic analyses based level in the early Pleistocene (1.45 Ma). Thus, although A. gym- on two- and six-marker datasets indicate that Aconitum subg. nandrum is a relatively ancient lineage in tribe Delphinieae, Gymnaconitum and Delphinium (sensu Jabbour & Renner, its population-level diversity originated more recently. When 2012a) grouped together (Figs. 3, 4). Thus, traditional Aconitum compared to the other three lineages, A. subg. Gymnaconitum is not monophyletic. When a phylogenetic tree conflicts with should therefore be regarded as a distinct taxon in tribe Del- the traditional concept of taxa (or an a priori hypothesis), it is phinieae. important to investigate if this concept lies within the range of Morphology. — Aconitum subg. Gymnaconitum differs possibilities supported by the data, which requires hypothesis markedly from Aconitum s.str. in having an annual or bi- testing. In our study, the SOWH tests support the exclusion of ennial habit (vs. perennial or pseudoannual), clawed sepals subgenus Gymnaconitum from Aconitum (Fig. 5). Based on (vs. clawless), large flabellate petaline labia (vs. small, not Bayesian relaxed-clock analyses, Jabbour & Renner (2012a) flabellate), 6–13 carpels (vs. 3–5), and subglobose seeds (vs. suggested that tribe Delphinieae rapidly diversified into four subpyramidal). The epidermal cells of the seeds of A. subg.

Tamura Jabbour & Renner Aconitella anthoroidea (1995) (2012) 51 0.96 Consolida persica 0.77 Consolida axilliflora 100 98 Consolida 1.00 1.00 Consolida ajacis 100 1.00 Consolida regalis 100 Consolida armeniaca 1.00 84 Delphinium delavayi Delphinium 1.00 100 Delphinium omeiense Delphinium Subg. Delphinastrum 57 1.00 Delphinium pseudohamatum 0.58 Delphinium tatsienense 100 100 Delphinium virgatum 0.02 1.00 73 1.00 Delphinium cossonianum Delphinium subg. Delphinium 1.00 Delphinium anthriscifolium Aconitum gymnandrum Aconitum subg. Gymnaconitum 98 Aconitum napellus 1.00 Aconitum pentheri 0.87 Aconitum baicalense 63 68 74 0.92 1.00 Aconitum ciliare 0.88 97 Aconitum delphinifolium Aconitum subg. Aconitum 1.00 100 72 Aconitum ferox 1.00 1.00 Aconitum pendulum Aconitum racemulosum Aconitum Aconitum barbatum 100 51 0.93 Aconitum septentrionale 100 1.00 97 Aconitum sinomontanum 1.00 98 1.00 99 0.90 Aconitum monticola 1.00 Aconitum subg. Lycoctonum 100 100 Aconitum longecassidatum 1.00 1.00 Aconitum alboviolaceum 57 Aconitum angustius 0.68 Aconitum scaposum 100 Delphinium staphisagria Delphinium subg. Staphisagria Staphisagria 1.00 Delphinium requienii Nigella damascena Outgroups Megaleranthis saniculifolia

Fig. 4. Phylogenetic relationships in tribe Delphinieae obtained from an ML analysis of the six-marker dataset. An ML phylogram is shown in the upper left hand corner. Numbers above and below branches are bootstrap values and Bayesian posterior probabilities (50%), respectively. Classifications by Tamura (1995) and Jabbour & Renner (2012a) are shown on the right.

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Gymnaconitum are round (Yang, 1990), whereas those in Based on above evidence, we raise A. subg. Gymnaconitum Aconitum s.str. are rectangular (Cappelletti & Poldini, 1984). to generic status, Gymnaconitum, below. Tribe Delphinieae Microstructural features of seed surfaces have been consid- now contains four genera: Staphisagria, Aconitum, Gymn ered to be less subject to environmental pressures than other aconitum, and Delphinium. morphological traits (Barthlott, 1984). Although A. subg. Gymnaconitum and Delphinium (sensu Jabbour & Renner, 2012a) form a clade (Figs. 3, 4), the two lineages differ in many TAXONOMY morphological characters, such as the upper sepal (no spur vs. long spur), petals (stalked vs. sessile), staminodes (absent Gymnaconitum (Stapf) Wei Wang & Z.D. Chen, stat. nov. ≡ vs. present), and carpels (many vs. several). A comparison of Aconitum sect. Gymnaconitum Stapf in Ann. Roy. Bot. morphological characters in tribe Delphinieae is shown in Gard. Calcutta 10: 178. 1905 ≡ Aconitum subg. Gymnaco- Table 2. Based on these characters, subgenus Gymnaconitum nitum (Stapf) Rapaics in Növényt. Közlem. 6: 139. 1907 should neither be part of traditional Aconitum nor be merged – Type: Gymnaconitum gymnandrum (Maxim.) Wei Wang with a broadly defined Delphinium. & Z.D. Chen. Karyology. — Based on plants collected from Qinghai, Gymnaconitum differs from Aconitum s.str. in having an Shang (1985) first reported that A. gymnandrum has a karyo- annual or biennial habit (vs. perennial or pseudoannual), cla- type formula of 2n = 16 = 12m + 4sm. Recently, Yuan (2006) wed sepals (vs. clawless), large flabellate petaline labia (vs. reported a karyotype formula of 2n = 16 = 6m + 10sm for small, not flabellate), 6–13 carpels (vs. 3–5), subglobose seeds this species from Sichuan. These two formulas are essentially (vs. subpyramidal), and round cells in the seed epidermis (vs. similar when carefully comparing the ratio of the two arms of rectangular). It is closely related to Delphinium sensu Jabbour each chromosome between two populations (Yuan, 2006). In & Renner, but differs markedly from the latter in the upper Aconitum s.str., the majority of species have only one or two sepal (no spur vs. long spur), petals (stalked vs. sessile), sta- pairs of m-chromosomes. Only A. chrysotrichum W.T. Wang minodes (absent vs. present), and number of carpels (many vs. and A. crassiflorum Hand.-Mazz. have karyotype formulas of several). 2n = 32 = 12m + 20sm. Some species also have st-chromosomes (Shang & Li, 1984; Verlaque & Aboucaya, 2001; Yang, 2001; Gymnaconitum gymnandrum (Maxim.) Wei Wang & Z.D. Yuan, 2006, and references therein). Staphisagria and Delphin- Chen, comb. nov. ≡ Aconitum gymnandrum Maxim. in ium (sensu Jabbour & Renner, 2012a) have a higher proportion Bull. Acad. Imp. Sci. Saint-Pétersbourg, ser. 3, 23(2): 308. of st- and/or t-chromosomes (Hong, 1986; Simon & al., 1995; 1877 – Lectotype (designated by Grubovskaya-Borodina Verlaque & Aboucaya, 2001; Yang, 2001; Bosch & al., 2002; in Grubov 2000: 189): China. Qinghai Prov., vicinity of Yuan, 2006). Thus, A. subg. Gymnaconitum has the most sym- Datong River, 14/26 Jul 1872, N.M. Przewalski s.n. (LE metric karyotype in tribe Delphinieae. [digital image!]).

Fig. 5. Histogram of the distribution of 100 replicates of the SOWH test of the monophyly of traditional Aconitum. Signifi- cance levels of 1% and 5% and the observed log-likelihood difference are shown in the chart.

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Table 2. Morphological comparison of Aconitum gymnandrum, Aconitum s.str., Staphisagria, and Delphinium (including Consolida; Jabbour & Renner, 2012a). Character Aconitum gymnandrum Aconitum s.str. Staphisagria Delphinium Habit a,b,c Annual or Perennial or Annual or Annual or biennial pseudoannual biennial perennial Sepalsa,b,d,e Clawed, Clawless, Clawless, Clawless, upper sepal not spurred upper sepal not spurred upper sepal spurred upper sepal spurred Petalsa,b,d,e Stalked, Stalked, Clawed, Sessile, lip large, flabellate lip not flabellate lip not flabellate lip not flabellate Staminodesb Absent Absent Present Present

Stigmaa,b Exposed Not exposed Not exposed Not exposed

No. of carpelsa,b 6–13 3 (6–8) or 3–5 (–9) 3 3 (4–10) or 1

Seed shapef,g Subglobose Subpyramidal Sector-sphaeroidal Subpyramidal to subglobose

Seed sizeb,f 1.5 mm 1.5–3.5 mm > 3.0 mm 1.0–3.5 mm

Seed surfacef,g Lamellae Lamellae or wings Reticulate Lamellae or wings

Epidermal cells of seedsf,g,h,i Round Rectangular Digitiform Rectangular a Tamura (1995); b W.T. Wang (1979); cJabbour & Renner (2012a); d Luo & Yang (2005); eJabbour & Renner (2012b); f Ilarslan & al. (1997); g Constantinidis & al. (2001); hYang (1990); i Cappelletti & Poldini (1984)

= Aconitum gymnandrum f. leucanthum W.T. Wang in Acta Muli, S.G. Wu 2891 (PE, KUN); Songpan, W.P. Fang 4013 Phytotax. Sin., Addit. 1: 97. 1965 – Holotype: China. Si- (NAS, IBSC, LBG); Litang, H.J. Wang 8257 (WUK); Garze, chuan Prov., Kangding, Xinduqiao, 21 Jul 1963, K.C. Kuan J.S. Yang 119 (PE, KUN); Shiqu, Y.H. Wu 29571 (HNWP); & W.T. Wang 869 (PE no. 00025217!). Sertar, Y.H. Wu 30808 (HNWP); Zoige, Sichuan Med. Exped. Note. – The type locality of A. gymnandrum was listed as 20340 (NAS); Maoxian, S. Jiang & J.C. Li 6835 (PE, KUN); “Chinae prov. Kansu” in the protologue. The provincial name Dawu, S. Jiang & C.J. Jin 2111 (PE, KUN); Ngawa, J.S. Yue Kansu is equivalent to Gansu. Nonetheless the type local- 64089 (NAS); Barkam, Z.R. Zhang 22623 (PE, KUN, NAS); ity, as inferred from the lectotype “China occidentalis. Terra Heishui, X. Li 73313 (PE, IBSC). Qinghai: Jigzhi, Y.H. Wu Tangutorum (prov. Kansu). Declivio australi jugi a fl. Tetung 26432 (HNWP); Ledu, Qinghai-Gansu Exped. 1716 (PE, meridiem versus”, is now situated in Qinghai Province and HNWP); Huzhu, Loess Exped. 5370 (WUK); Gonghe, has not been included within Gansu Province since the early X.S. Wang 149 (HNWP); Xinghai, Z.B. Wang 20280 (WUK, twentieth century. HNWP); Gangca, S.X. Wang 118 (HNWP); Tongren, Y.H. Wu Icon. – Institute of Botany, Chinese Academy of Sciences 32394 (HNWP); Tongde, Y.H. WU 21482 (HNWP); Nangqen, (1972: fig. 1396); W.T. Wang (1979: fig. 77, 1–3). Y.C. Yang 01188 (PE, HNWP); Jainca, L.H. Tang & L.N. Sun Distribution and habitat. – China (Xizang [Tibet], W 733 (HNWP); Pingan, Y.H. Wu & al. 4218 (HNWP); Xunhua, Sichuan, Qinghai, and S Gansu). Alpine grassy and sandy Y.H. Wu 28492 (HNWP); Qumarlêb, R.F. Huang 156 (HNWP); slopes, grasslands; 1550–3800 m. Zadoi, S.W. Liu 302 (HNWP, WUK); Golog, W.Y. Wang 27291 Representative specimens examined. – CHINA. Xizang (HNWP); Minhe, T.N. Ho 572 (HNWP); Zekog, S.W. Liu (Tibet): Lasa, J.S. Yang 53 (KUN); Lhünze, Qinghai-Tibet & D.S. Luo 1584 (HNWP),B.Z. Guo 25756 (HNWP); Maqen, Exped. 2902 (PE); Tingri, R.F. Huang CG89-768 (HNWP); T.N. Ho & al. 436 (PE, HNWP); Baima, Y.H. Wu 26078 Jomda, Qinghai-Tibet Exped. 2006 (PE, KUN); Ngamring, (HNWP); Gade, Y.H. Wu 25729 (HNWP); Qilian, T.N. Ho 3223 Qinghai-Tibet 7312 (PE, KUN); Bomê, Qinghai-Tibet Sci- (HNWP); Xining, Z.H. Zhang & al. 0776 (HNWP); Guinan, ence Exped. 12582 (PE); Qamdo, Biological Institute Xizang Y.H. Wu 21003 (HNWP); Menyuan, T.S. Ying & al. 26931 (PE, Exped. 2223 (HNWP); Riwuqê, Qinghai-Tibet Exped. 12971 KUN). Gansu: Lintan, Yellow River Exped. 3303 (PE, NAS, (PE, KUN); Konjo, Qinghai-Tibet Exped. 12582 (PE, KUN, KUN); Lanzhou, Y.Q. He 4378 (WUK); Tianzhu, Y.Q. He 4554 HNWP); Nagqu, D.D. Tao 10843 (PE, KUN); Cona, C.Y. Wu (WUK); Kangle, Taohe Exped. 95 (KUN); Jingtai, Z.Y. Yu & al. 75-911 (PE, HNWP, KUN); Yadong, Qinghai-Tibet & Y.P. Xu 3127 (WUK); Yuzhong, Yellow RiverExped. 3031 Exped. 74-2579 (PE). Sichuan: Xiangcheng, Qinghai-Tibet (PE, WUK); Maqu, Y.H. Wu 31750 (HNWP); Zhugqu, B.Z. Guo Exped. 4856 (KUN); Kangding, S. Jiang 6412 (PE, KUN); 5606 (WUK).

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ACKNOWLEDGEMENTS Jabbour, F. & Renner, S. 2011a. Consolida and Aconitella are an annual clade of Delphinium (Ranunculaceae) that diversified in the Mediterranean basin and the Irano-Turanian region. Taxon We sincerely thank Prof. Pei-Gen Xiao and An-Ming Lu for their 60: 1029–1040. helpful suggestions; Dr. Hong-Lei Li and Cai-Fei Zhang for sequence Jabbour, F. & Renner, S. 2011b. Resurrection of the genus Staphis- and literature collections; Dr. Lily R. Lewis for her carefully read- agria J. Hill, sister to all the other Delphinieae (Ranunculaceae). ing the manuscript and helpful suggestions; the curators of HNWP, PhytoKeys 7: 21–26. http://dx.doi.org/10.3897/phytokeys.7.2010 KUN, NAS, PE and WUK for accessing to their specimens; Dr. Jabbour, F. & Renner, S. 2012a. A phylogeny of Delphinieae (Ranun- Irina Illarionova (LE) for providing digital type images. 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Gymnaconitum, a new genus of Ranunculaceae endemic to the Qinghai-Tibetan Plateau Wei Wang, Yang Liu, Sheng-Xiang Yu, Tian-Gang Gao & Zhi-Duan Chen

Taxon : – TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

Consolida pubescens Consolida regalis Fig. S1. Phylogram of 179 acces- Consolida persica Aconitella aconiti Aconitella thirkeana sions obtained from ML analysis Aconitella stenocarpa Consolida aucheri Consolida kandaharica of the trnL-F data. Numbers at Consolida flava Consolida camptocarpa 95 Consolida kabuliana the nodes are bootstrap percent- Consolida stocksiana Consolida songorica Aconitella barbata ages (> 70%). Classification of Consolida mauritanica Consolida Consolida orientalis Aconitella hohenackeri Tamura (1995) is shown on the 79 Aconitella anthoroidea Consolida hellespontica right. Consolida glandulosa Consolida hispanica Consolida axilliflora Consolida ajacis Consolida uechtritziana 100 Consolida lineolata 92 Consolida armeniaca Consolida olopetala Delphinium grandiflorum Delphinium tatsienense Delphinium cheilanthum Delphinium sparsiflorum Delphinium kansuense Delphinium middendorffii Delphinium brachycentrum Delphinium kamaonense 81 Delphinium menziesii Delphinium depauperatum Delphinium stachydeum Delphinium andersonii Delphinium bicolor Delphinium barbeyi Delphinium exaltatum Delphinium nuttallianum Delphinium multiplex Delphinium scaposum 86 Delphinium viridescens Delphinium glaucescens Delphinium trolliifolium Delphinium nudicaule Delphinium parryi Delphinium gypsophilum Delphinium decorum Delphinium californicum Delphinium polycladon Delphinium recurvatum Delphinium hesperium Delphinium parishii Delphinium patens Delphinium bakeri Delphinium cardinale 83 Delphinium hutchinsonae Delphinium caeruleum Delphinium viscosum Delphinium tangkulaense Delphinium delavayi Delphinium pylzowii Delphinium potaninii Delphinium bonvalotii Delphinium crispulum Delphinium elbursense Delphinium dolichostachyum Delphinium flexuosum Delphinium oreophilum 75 Delphinium cashmerianum 98 Delphinium minjanense Delphinium afghanicum Delphinium bulleyanum Delphinium subg. Delphinastrum 75 Delphinium montanum Delphinium dubium 72 Delphinium cuneatum Delphinium oxysepalum 83 Delphinium elatum Delphinium crassifolium Delphinium maackianum 75 Delphinium pyramidale Delphinium muscosum Delphinium glaciale Delphinium ochotense Delphinium beesianum Delphinium gyalanum Delphinium carduchorum Delphinium szowitsianum Delphinium aquilegifolium Delphinium cyphoplectrum Delphinium tuberosum Delphinium albocoeruleum Delphinium biternatum Delphinium griseum Delphinium denudatum Delphinium suave Delphinium wendelboi Delphinium saniculifolium Delphinium semibarbatum Delphinium macrostachyum Delphinium fissum Delphinium freynii Delphinium kurdicum Delphinium kohatense 87 Delphinium dasycaulon Delphinium macrocentron Delphinium leroyi Delphinium incisum 72 Delphinium pentagynum Delphinium nevadense Delphinium sylvaticum Delphinium emarginatum Delphinium sutchuenense Delphinium tsarongense Delphinium orthocentrum Delphinium pachycentrum 70 Delphinium trichophorum Delphinium oxycentrum Delphinium thibeticum Delphinium anthriscifolium Delphinium cossonianum Delphinium macropetalum Delphinium verdunense Delphinium gracile Delphinium hirschfeldianum 92 Delphinium halteratum 100 Delphinium balcanicum Delphinium subg. Delphinium Delphinium peregrinum 96 Delphinium obcordatum Delphinium nanum 98 Delphinium venulosum Delphinium virgatum Delphinium balansae Delphinium favargeri Aconitum gymnandrum Aconitum subg. Gymnaconitum Aconitum pendulum Aconitum nagarum Aconitum sessiliflorum Aconitum delphinifolium Aconitum baicalense Aconitum pentheri Aconitum chienningense Aconitum forrestii Aconitum pulchellum Aconitum hemsleyanum Aconitum ferox Aconitum liljestrandii Aconitum franchetii Aconitum ciliare Aconitum kitadakense Aconitum subg. Aconitum 100 Aconitum paskoi Aconitum yamazakii + Aconitum sanyoense Aconitum senanense Aconitum subg. Lycoctonum Aconitum decipiens Aconitum villosum Aconitum yuparense Aconitum nipponicum Aconitum tanguticum Aconitum racemulosum Aconitum violaceum 99 Aconitum chrysotrichum Aconitum crassiflorum Aconitum sinomontanum 71 100 Aconitum scaposum Aconitum gigas Aconitum finetianum Aconitum septentrionale Aconitum lycoctonum Aconitum iidemontanum 100 Delphinium pictum Delphinium requienii Delphinium subg. Staphisagria Delphinium staphisagria Nigella damascena Megaleranthis saniculifolia 0.01

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Fig. S2. Phylogram of 179 acces- Consolida stocksiana Consolida songorica 0.98 Consolida hellespontica sions obtained from BI analysis Consolida glandulosa Consolida uechtritziana Consolida axilliflora of the trnL-F data. Numbers at Consolida ajacis Aconitella thirkeana Aconitella stenocarpa the nodes are Bayesian posterior Consolida orientalis Consolida mauritanica 1.0 Aconitella barbata probabilities (> 0.95). Classifica- Aconitella hohenackeri Aconitella anthoroidea Consolida regalis Consolida tion of Tamura (1995) is shown on Consolida pubescens Consolida persica Consolida kandaharica the right. 0.99 Consolida kabuliana Consolida hispanica Consolida flava Consolida camptocarpa Consolida aucheri Aconitella aconiti 1.0 Consolida lineolata Consolida armeniaca Consolida olopetala Delphinium beesianum Delphinium bonvalotii Delphinium caeruleum Delphinium dolichostachyum Delphinium flexuosum Delphinium glaciale Delphinium gyalanum Delphinium ochotense Delphinium tangkulaense Delphinium viscosum 0.99 Delphinium afghanicum Delphinium minjanense Delphinium cashmerianum 0.99 Delphinium oreophilum Delphinium incisum Delphinium kohatense Delphinium albocoeruleum Delphinium biternatum Delphinium denudatum 0.99 Delphinium griseum Delphinium saniculifolium Delphinium semibarbatum Delphinium suave Delphinium wendelboi 0.97 Delphinium carduchorum Delphinium aquilegifolium Delphinium cyphoplectrum 0.99 Delphinium szowitsianum Delphinium tuberosum 0.99 Delphinium dasycaulon Delphinium leroyi Delphinium macrocentron Delphinium nevadense 1.0 Delphinium pentagynum 0.98 Delphinium emarginatum Delphinium sylvaticum 1.00 1.0 Delphinium freynii Delphinium kurdicum Delphinium fissum Delphinium macrostachyum Delphinium bulleyanum Delphinium dubium Delphinium montanum Delphinium cuneatum Delphinium elatum 0.97 Delphinium oxysepalum Delphinium crassifolium Delphinium maackianum Delphinium crispulum Delphinium elbursense Delphinium Subg. Delphinastrum Delphinium delavayi Delphinium potaninii Delphinium pylzowii Delphinium muscosum Delphinium pyramidale Delphinium cheilanthum Delphinium grandiflorum Delphinium kansuense 1.00 Delphinium middendorffii Delphinium sparsiflorum Delphinium tatsienense Delphinium multiplex Delphinium nuttallianum Delphinium scaposum Delphinium andersonii 0.96 Delphinium bicolor Delphinium depauperatum Delphinium menziesii Delphinium stachydeum Delphinium bakeri Delphinium californicum Delphinium cardinale 1.0 Delphinium decorum 0.95 Delphinium gypsophilum Delphinium hesperium 0.97 Delphinium hutchinsonae Delphinium nudicaule Delphinium parishii Delphinium parryi Delphinium patens Delphinium polycladon Delphinium recurvatum Delphinium trolliifolium 0.98 Delphinium barbeyi Delphinium exaltatum 0.98 Delphinium glaucescens Delphinium viridescens Delphinium brachycentrum Delphinium kamaonense Delphinium oxycentrum Delphinium thibeticum 0.97 Delphinium sutchuenense Delphinium pachycentrum 0.99 Delphinium trichophorum 0.99 Delphinium orthocentrum 0.99 Delphinium tsarongense 0.96 Delphinium virgatum Delphinium venulosum 1.0 Delphinium obcordatum Delphinium nanum 1.0 Delphinium peregrinum Delphinium halteratum Delphinium balcanicum Delphinium verdunense 0.96 1.00 Delphinium macropetalum Delphinium subg. Delphinium Delphinium hirschfeldianum Delphinium gracile Delphinium favargeri Delphinium cossonianum Delphinium balansae Delphinium anthriscifolium Aconitum gymnandrum Aconitum subg. Gymnaconitum Aconitum baicalense Aconitum chienningense Aconitum ciliare Aconitum decipiens Aconitum delphinifolium Aconitum ferox Aconitum forrestii Aconitum franchetii Aconitum hemsleyanum Aconitum kitadakense Aconitum liljestrandii Aconitum nagarum Aconitum nipponicum Aconitum paskoi 1.00 Aconitum pendulum Aconitum pentheri Aconitum pulchellum Aconitum subg. Aconitum Aconitum racemulosum Aconitum sanyoense Aconitum senanense + Aconitum sessiliflorum Aconitum subg. Lycoctonum Aconitum tanguticum Aconitum villosum Aconitum violaceum 0.97 Aconitum yamazakii Aconitum yuparense Aconitum lycoctonum Aconitum scaposum 1.0 Aconitum septentrionale 1.0 Aconitum chrysotrichum Aconitum crassiflorum Aconitum sinomontanum 0.96 Aconitum finetianum Aconitum gigas Aconitum iidemontanum 1.00 Delphinium pictum Delphinium requienii Delphinium subg. Staphisagria Delphinium staphisagria Nigella damascena Megaleranthis saniculifolia 0.5

Version of Record. S2 TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

Fig. S3. Phylogram of 234 acces- continued sions obtained from ML analysis next page Delphinium halteratum of the ITS data. Numbers at the Delphinium balcanicum Delphinium balansae nodes are bootstrap percentages Delphinium cossonianum 96 Delphinium obcordatum (> 70%). Classification of Tamura 76 Delphinium verdunense Delphinium favargeri Delphinium subg. Delphinium (1995) is shown on the right. Delphinium gracile 94 Delphinium venulosum 100 Delphinium macropetalum Delphinium peregrinum Delphinium hirschfeldianum Delphinium virgatum Delphinium anthriscifolium Aconitum campylorrhynchum Aconitum bulleyanum Aconitum georgei Aconitum bulbilliferum Aconitum piepunense Aconitum legendrei Aconitum forrestii Aconitum vilmorinianum Aconitum contortum 93 Aconitum transsectum Aconitum delavayi Aconitum stapfianum Aconitum hemsleyanum 86 Aconitum fengii 90 Aconitum pilopetalum Aconitum tongolense Aconitum refractum Aconitum chienningense Aconitum tuguancunense Aconitum tatsienense Aconitum liljestrandii Aconitum franchetii Aconitum tsaii Aconitum episcopale Aconitum nemorum Aconitum decipiens Aconitum karakolicum Aconitum cannabifolium 96 83 Aconitum napellus 96 Aconitum variegatum Aconitum pentheri 78 Aconitum paskoi Aconitum baicalense Aconitum monanthum 94 Aconitum apoiense Aconitum yamazakii Aconitum yuparense Aconitum chiisanense Aconitum sanyoense Aconitum iidemontanum Aconitum subg. Aconitum Aconitum austrokoreense Aconitum nipponicum Aconitum hakusanense Aconitum senanense Aconitum kitadakense Aconitum villosum Aconitum ciliare Aconitum jaluense Aconitum napiforme Aconitum bartlettii Aconitum yezoense Aconitum volubile Aconitum carmichaelii Aconitum kusnezoffii 94 Aconitum delphinifolium Aconitum columbianum Aconitum sungpanense Aconitum pendulum Aconitum balfourii 99 Aconitum palmatum Aconitum ferox Aconitum pulchellum Aconitum liangshanicum Aconitum nagarum 84 Aconitum hamatipetalum 81Aconitum sessiliflorum Aconitum flavum Aconitum violaceum Aconitum tanguticum 86Aconitum huiliense Aconitum brachypodum Aconitum spiripetalum Aconitum heterophyllum Aconitum brunneum 86 Aconitum zeravschanicum Aconitum anthora Aconitum racemulosum Aconitum polycarpum 80 Aconitum lasiostomum Aconitum lycoctonum 100 Aconitum orientale 100 Aconitum septentrionale Aconitum moldavicum Aconitum monticola 90 75 Aconitum sinomontanum Aconitum shennongjiaense 100Aconitum longecassidatum 99 Aconitum finetianum Aconitum subg. Lycoctonum Aconitum alboviolaceum Aconitum umbrosum 91 Aconitum gigas Aconitum barbatum Aconitum scaposum 100 100Aconitum chrysotrichum Aconitum crassiflorum Aconitum angustius Aconitum gymnandrum Aconitum subg. Gymnaconitum 100 Delphinium staphisagria Delphinium requienii Delphinium subg. Staphisagria Nigella damascena Megaleranthis saniculifolia 0.1

Version of Record. S3 TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

100 Aconitella aconiti 96 Aconitella thirkeana Aconitella stenocarpa 100 Aconitella hohenackeri Aconitella anthoroidea 99 Consolida hispanica Consolida orientalis 74 99 Consolida ajacis Consolida uechtritziana 72 Consolida axilliflora 99 Consolida songorica Consolida camptocarpa 100 Consolida kandaharica Consolida aucheri Consolida 99 Consolida stocksiana 100 Consolida kabuliana Consolida flava 71 Consolida persica Consolida regalis Aconitella barbata 100 100 Consolida glandulosa 96 Consolida hellespontica 97 Consolida pubescens Consolida mauritanica 72 Consolida olopetala 100 Consolida lineolata Consolida armeniaca Delphinium oxycentrum Delphinium trichophorum Delphinium forrestii Delphinium viscosum Delphinium pentagynum Delphinium dolichostachyum Delphinium thibeticum Delphinium parryi Delphinium gypsophilum Delphinium cardinale Delphinium nuttallii Delphinium glaucum Delphinium sutherlandii Delphinium bicolor Delphinium glareosum Delphinium xantholeucum Delphinium glaucescens Delphinium brachycentrum Delphinium patens 84 Delphinium depauperatum Delphinium nuttallianum Delphinium lineapetalum Delphinium menziesii Delphinium hutchinsoniae Delphinium californicum Delphinium inopinum Delphinium viridescens Delphinium trolliifolium Delphinium antoninum Delphinium nudicaule Delphinium bakeri Delphinium stachydeum Delphinium andersonii Delphinium gracilentum Delphinium parishii Delphinium purpusii Delphinium variegatum Delphinium barbeyi Delphinium scaposum Delphinium wislizeni 100 95 Delphinium bicornutum Delphinium calcarequitis 96 Delphinium subscandens Delphinium viride 72 Delphinium sapellonis 85 Delphinium valens 88 Delphinium alpestre Delphinium subg. Delphinastrum Delphinium novomexicanum Delphinium ramosum Delphinium multiplex Delphinium distichum Delphinium hansenii Delphinium hesperium 94 Delphinium andesicola Delphinium geraniifolium Delphinium ulignosum Delphinium polycladon Delphinium geyeri 98 Delphinium wootonii Delphinium carolinianum Delphinium madrense Delphinium recurvatum Delphinium luteum Delphinium umbraculorum Delphinium decorum Delphinium newtonianum 78 Delphinium alabamicum 84 Delphinium tricorne Delphinium treleasei 71 Delphinium oxysepalum Delphinium spirocentrum Delphinium montanum Delphinium tatsienense Delphinium orthocentrum 79Delphinium grandiflorum Delphinium kamaonense Delphinium ceratophorum Delphinium potaninii 79 Delphinium pseudohamatum Delphinium dolichocentroides Delphinium pycnocentrum Delphinium pachycentrum Delphinium sutchuenense Delphinium pyramidale Delphinium muscosum Delphinium smithianum Delphinium sparsiflorum Delphinium batangense 100 Delphinium taliense Delphinium muliense Delphinium delavayi continued Delphinium omeiense previous page

0.1

Version of Record. S4 TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

Fig. S4. Phylogram of 234 acces- continued sions obtained from BI analysis next page Delphinium peregrinum of the ITS data. Numbers at the Delphinium hirschfeldianum Delphinium verdunense nodes are Bayesian posterior 0.99 Delphinium favargeri Delphinium obcordatum probabilities (> 0.95). Classifica- Delphinium cossonianum Delphinium gracile Delphinium subg. Delphinium tion of Tamura (1995) is shown on Delphinium halteratum 0.98 1.00 Delphinium balcanicum the right. Delphinium venulosum Delphinium macropetalum Delphinium balansae Delphinium virgatum Delphinium anthriscifolium Aconitum gymnandrum Aconitum subg. Gymnaconitum Aconitum tsaii 0.99 Aconitum nemorum Aconitum karakolicum Aconitum episcopale Aconitum decipiens Aconitum heterophyllum Aconitum brunneum Aconitum zeravschanicum Aconitum tanguticum Aconitum spiripetalum Aconitum sessiliflorum Aconitum flavum Aconitum violaceum 1.0 Aconitum palmatum Aconitum ferox Aconitum pulchellum Aconitum nagarum Aconitum liangshanicum Aconitum hamatipetalum 0.99 Aconitum huiliense Aconitum brachypodum Aconitum pendulum Aconitum balfourii 0.97 Aconitum tatsienense Aconitum liljestrandii Aconitum franchetii 1.0Aconitum delphinifolium Aconitum columbianum Aconitum cannabifolium 1.0 Aconitum tongolense 0.99 Aconitum pilopetalum Aconitum stapfianum Aconitum delavayi Aconitum hemsleyanum Aconitum fengii Aconitum contortum Aconitum vilmorinianum Aconitum tuguancunense Aconitum transsectum 1.0 Aconitum refractum Aconitum subg. Aconitum Aconitum piepunense Aconitum legendrei Aconitum georgei Aconitum forrestii Aconitum chienningense Aconitum campylorrhynchum Aconitum bulleyanum Aconitum bulbilliferum Aconitum senanense Aconitum kitadakense 0.96 Aconitum nipponicum 1.00 Aconitum hakusanense 0.96Aconitum jaluense Aconitum ciliare Aconitum yezoense Aconitum bartlettii Aconitum volubile Aconitum napiforme Aconitum kusnezoffii Aconitum carmichaelii Aconitum villosum Aconitum variegatum 1.0 Aconitum napellus 1.00 0.99 Aconitum pentheri Aconitum paskoi 1.0 Aconitum baicalense 0.98Aconitum yamazakii Aconitum apoiense Aconitum yuparense Aconitum sanyoense Aconitum iidemontanum 1.00 Aconitum chiisanense Aconitum monanthum Aconitum austrokoreense Aconitum sungpanense Aconitum racemulosum Aconitum polycarpum Aconitum anthora 0.98 Aconitum sinomontanum Aconitum monticola Aconitum lycoctonum 0.99 Aconitum lasiostomum Aconitum shennongjiaense Aconitum septentrionale Aconitum orientale 1.0 Aconitum moldavicum Aconitum umbrosum 0.98 Aconitum gigas Aconitum subg. Lycoctonum Aconitum barbatum Aconitum longecassidatum 1.0 Aconitum finetianum 1.00 Aconitum alboviolaceum Aconitum scaposum Aconitum angustius Aconitum chrysotrichum 1.0 Aconitum crassiflorum 1.00 Delphinium staphisagria Delphinium requienii Delphinium subg. Staphisagria Nigella damascena Megaleranthis saniculifolia 0.2

Version of Record. S5 TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

1.0 Aconitella thirkeana 1.0 Aconitella aconiti Aconitella stenocarpa 1.0 Aconitella hohenackeri Aconitella anthoroidea 0.99 Consolida pubescens 1.0 Consolida mauritanica 1.0 Consolida hellespontica Consolida glandulosa Aconitella barbata 0.99 Consolida persica Consolida flava 1.0 0.99 Consolida stocksiana Consolida kabuliana Consolida 1.0 1.0 Consolida songorica 0.95 Consolida camptocarpa 1.0 Consolida kandaharica Consolida aucheri 1.0 Consolida orientalis 1.00 Consolida hispanica 1.0 Consolida uechtritziana Consolida ajacis Consolida axilliflora Consolida regalis Consolida lineolata 1.0 Consolida armeniaca Consolida olopetala 1.0 Delphinium kamaonense Delphinium grandiflorum 0.98 Delphinium pseudohamatum Delphinium dolichocentroides Delphinium pycnocentrum 0.99 Delphinium sparsiflorum Delphinium smithianum Delphinium pyramidale 0.99 Delphinium muscosum Delphinium batangense Delphinium omeiense Delphinium muliense Delphinium delavayi Delphinium purpusii Delphinium parishii Delphinium hesperium Delphinium hansenii 1.0 Delphinium nuttallianum Delphinium lineapetalum 0.97 Delphinium depauperatum Delphinium menziesii 1.0 Delphinium recurvatum Delphinium luteum 1.0 Delphinium umbraculorum Delphinium decorum Delphinium gypsophilum Delphinium parryi 1.0 Delphinium wootonii Delphinium madrense Delphinium carolinianum 0.96Delphinium hutchinsoniae Delphinium californicum Delphinium subscandens 1.0 Delphinium calcarequitis 1.0 Delphinium viride Delphinium wislizeni Delphinium bicornutum Delphinium variegatum Delphinium scaposum 1.00 Delphinium barbeyi 0.95 Delphinium trolliifolium Delphinium antoninum Delphinium nudicaule Delphinium bakeri 1.0 Delphinium geraniifolium Delphinium andesicola Delphinium valens Delphinium subg. Delphinastrum 1.0 0.99 Delphinium sapellonis Delphinium novomexicanum Delphinium alpestre Delphinium ramosum 0.95 Delphinium tricorne Delphinium alabamicum Delphinium treleasei Delphinium newtonianum Delphinium xantholeucum Delphinium viridescens Delphinium ulignosum Delphinium sutherlandii 0.99 Delphinium stachydeum Delphinium polycladon Delphinium patens Delphinium nuttallii Delphinium multiplex Delphinium inopinum Delphinium gracilentum Delphinium glaucum Delphinium glaucescens Delphinium glareosum Delphinium geyeri Delphinium distichum Delphinium cardinale 0.99 Delphinium brachycentrum Delphinium bicolor Delphinium andersonii Delphinium oxysepalum Delphinium spirocentrum Delphinium montanum Delphinium tatsienense Delphinium thibeticum Delphinium taliense Delphinium sutchuenense 0.97 Delphinium potaninii Delphinium pachycentrum Delphinium orthocentrum Delphinium ceratophorum 1.00 Delphinium dolichostachyum Delphinium pentagynum Delphinium trichophorum Delphinium oxycentrum 0.2 Delphinium forrestii continued Delphinium viscosum previous page

Version of Record. S6 TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

Fig. S5. Phylogram of 287 acces- continued sions obtained from ML analysis next page Delphinium balansae of the two-marker dataset. Num- Delphinium hirschfeldianum 74 Delphinium halteratum bers at the nodes are bootstrap 89 Delphinium balcanicum Delphinium peregrinum Delphinium gracile percentages (>70%). Classifica- Delphinium favargeri tion of Tamura (1995) is shown on 97 Delphinium cossonianum Delphinium subg. Delphinium 99 70 Delphinium verdunense 100 Delphinium obcordatum the right. 100 Delphinium nanum Delphinium macropetalum 65 Delphinium venulosum Delphinium virgatum Delphinium anthriscifolium Aconitum gymnandrum Aconitum subg. Gymnaconitum Aconitum zeravschanicum Aconitum brunneum Aconitum heterophyllum Aconitum spiripetalum Aconitum tanguticum Aconitum violaceum Aconitum flavum 76 Aconitum sessiliflorum Aconitum balfourii Aconitum pendulum Aconitum hamatipetalum Aconitum pulchellum Aconitum palmatum 98 Aconitum ferox Aconitum nagarum Aconitum liangshanicum Aconitum iidemontanum Aconitum sanyoense Aconitum chiisanense Aconitum apoiense 91 Aconitum yamazakii Aconitum yuparense Aconitum austrokoreense Aconitum monanthum Aconitum baicalense Aconitum paskoi 85 Aconitum pentheri 92 96 Aconitum variegatum 91 Aconitum napellus Aconitum legendrei 96 Aconitum transsectum Aconitum piepunense Aconitum bulleyanum Aconitum forrestii Aconitum bulbilliferum

4 Aconitum georgei Aconitum campylorrhynchum 80 Aconitum vilmorinianum Aconitum refractum 88 Aconitum pilopetalum Aconitum subg. Aconitum Aconitum tongolense Aconitum fengii Aconitum hemsleyanum 89Aconitum stapfianum Aconitum delavayi Aconitum contortum Aconitum chienningense Aconitum tuguancunense Aconitum sungpanense Aconitum franchetii Aconitum tatsienense 78 Aconitum liljestrandii 96 Aconitum columbianum Aconitum delphinifolium Aconitum ciliare Aconitum jaluense Aconitum yezoense Aconitum bartlettii 98 Aconitum napiforme Aconitum kusnezoffii Aconitum volubile Aconitum carmichaelii Aconitum nipponicum 100 Aconitum hakusanense Aconitum villosum Aconitum kitadakense 82 Aconitum senanense Aconitum cannabifolium Aconitum karakolicum Aconitum decipiens 70 Aconitum nemorum Aconitum tsaii Aconitum episcopale 99 Aconitum huiliense 93 Aconitum brachypodum Aconitum racemulosum Aconitum anthora Aconitum polycarpum 74 Aconitum lasiostomum Aconitum lycoctonum Aconitum orientale 97 Aconitum moldavicum 76Aconitum sinomontanum Aconitum monticola Aconitum shennongjiaense 99 Aconitum septentrionale Aconitum barbatum 91 Aconitum gigas Aconitum subg. Lycoctonum Aconitum umbrosum 100 74 Aconitum alboviolaceum Aconitum longecassidatum 100 Aconitum finetianum Aconitum scaposum Aconitum angustius Aconitum chrysotrichum 100 Aconitum crassiflorum 100 Delphinium staphisagria Delphinium pictum Delphinium requienii Delphinium subg. Staphisagria Nigella damascena Megaleranthis saniculifolia 0.02

Version of Record. S7 TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

99 97 Aconitella thirkeana Aconitella aconiti 100 Aconitella stenocarpa Aconitella anthoroidea Aconitella hohenackeri 100 Consolida uechtritziana 77 Consolida ajacis 86 99 Consolida hispanica Consolida orientalis Consolida axilliflora 98 Consolida kabuliana Consolida stocksiana 100 Consolida camptocarpa Consolida songorica 91 100 100 Consolida kandaharica Consolida Consolida aucheri Consolida flava Consolida persica Consolida regalis Aconitella barbata 100 97 Consolida pubescens 97 Consolida mauritanica 100 Consolida hellespontica Consolida glandulosa 100 Consolida armeniaca Consolida lineolata Consolida olopetala Delphinium caeruleum Delphinium tangkulaense Delphinium szowitsianum Delphinium aquilegifolium Delphinium cyphoplectrum Delphinium tuberosum Delphinium carduchorum Delphinium sylvaticum Delphinium emarginatum Delphinium pentagynum Delphinium nevadense Delphinium dasycaulon Delphinium macrocentron Delphinium leroyi 77 Delphinium macrostachyum Delphinium fissum Delphinium freynii Delphinium kurdicum Delphinium biternatum Delphinium saniculifolium Delphinium denudatum Delphinium griseum Delphinium wendelboi Delphinium suave Delphinium semibarbatum Delphinium albocoeruleum Delphinium kohatense Delphinium incisum Delphinium taliense Delphinium muliense 77 Delphinium muscosum Delphinium pyramidale Delphinium smithianum Delphinium dolichostachyum Delphinium flexuosum Delphinium afghanicum Delphinium minjanense Delphinium oreophilum Delphinium cashmerianum Delphinium elbursense Delphinium crispulum Delphinium maackianum Delphinium crassifolium Delphinium bonvalotii Delphinium beesianum Delphinium glaciale Delphinium gyalanum 85 Delphinium pseudohamatum Delphinium dolichocentroides Delphinium pycnocentrum Delphinium potaninii Delphinium pylzowii Delphinium delavayi Delphinium ochotense 76 Delphinium grandiflorum Delphinium kamaonense Delphinium kansuense Delphinium spirocentrum Delphinium ramosum 84 77 Delphinium valens Delphinium sapellonis Delphinium alpestre Delphinium subg. Delphinastrum 99 88 Delphinium novomexicanum Delphinium distichum Delphinium multiplex 77 Delphinium wislizeni 97 Delphinium bicornutum 70 Delphinium calcarequitis Delphinium subscandens 95 Delphinium viride Delphinium glareosum Delphinium xantholeucum Delphinium lineapetalum Delphinium nuttallianum Delphinium depauperatum Delphinium menziesii Delphinium sutherlandii Delphinium bicolor Delphinium stachydeum Delphinium gracilentum Delphinium inopinum Delphinium andersonii Delphinium viridescens Delphinium glaucescens Delphinium geyeri Delphinium polycladon Delphinium purpusii Delphinium parishii Delphinium variegatum Delphinium exaltatum Delphinium barbeyi Delphinium scaposum Delphinium hesperium Delphinium hansenii Delphinium ulignosum 94 Delphinium andesicola Delphinium geraniifolium Delphinium parryi Delphinium gypsophilum Delphinium luteum Delphinium recurvatum Delphinium umbraculorum Delphinium decorum Delphinium cardinale Delphinium nudicaule Delphinium bakeri Delphinium trolliifolium Delphinium antoninum Delphinium patens Delphinium hutchinsonae 73 Delphinium californicum Delphinium nuttallii Delphinium glaucum 100 Delphinium madrense Delphinium carolinianum Delphinium wootonii Delphinium brachycentrum Delphinium treleasei Delphinium newtonianum Delphinium tricorne 76 Delphinium alabamicum Delphinium oxysepalum Delphinium elatum Delphinium cuneatum Delphinium bulleyanum Delphinium montanum Delphinium dubium Delphinium tatsienense Delphinium omeiense Delphinium middendorffii Delphinium cheilanthum Delphinium sparsiflorum Delphinium batangense Delphinium ceratophorum Delphinium tsarongense 99 Delphinium orthocentrum Delphinium pachycentrum Delphinium sutchuenense Delphinium thibeticum 0.02 Delphinium oxycentrum Delphinium trichophorum Delphinium forrestii continued Delphinium viscosum previous page Version of Record. S8 TAXON 62 (4) • August 2013 Electr. Suppl. to: Wang & al. • Gymnaconitum, a new genus of Ranunculaceae

Fig. S6. Phylogram of 287 acces- continued sions obtained from BI analysis of next page 1.0 Delphinium obcordatum the two-marker dataset. Num- Delphinium nanum 1.0 Delphinium favargeri bers at the nodes are Bayesian 0.98 Delphinium cossonianum Delphinium verdunense posterior probabilities (> 0.95). Delphinium gracile 0.97 Delphinium halteratum 0.99 1.0 Delphinium balcanicum Delphinium subg. Delphinium Classification of Tamura (1995) is Delphinium peregrinum 1.00 Delphinium macropetalum Delphinium hirschfeldianum shown on the right. 1.00 0.97 Delphinium balansae Delphinium venulosum Delphinium virgatum Delphinium anthriscifolium Aconitum gymnandrum Aconitum subg. Gymnaconitum Aconitum tanguticum Aconitum spiripetalum Aconitum heterophyllum Aconitum brunneum Aconitum zeravschanicum Aconitum sessiliflorum Aconitum flavum Aconitum violaceum 1.0 Aconitum palmatum Aconitum ferox Aconitum pulchellum Aconitum nagarum Aconitum liangshanicum Aconitum hamatipetalum 0.99Aconitum huiliense Aconitum brachypodum Aconitum pendulum Aconitum balfourii Aconitum tsaii 0.98 Aconitum nemorum Aconitum karakolicum Aconitum episcopale Aconitum decipiens 0.98 Aconitum tatsienense Aconitum liljestrandii Aconitum franchetii 1.0 Aconitum delphinifolium Aconitum columbianum Aconitum cannabifolium 1.0 Aconitum tongolense Aconitum pilopetalum Aconitum stapfianum Aconitum delavayi 1.00 Aconitum hemsleyanum Aconitum fengii Aconitum contortum Aconitum vilmorinianum Aconitum tuguancunense Aconitum transsectum 1.0 Aconitum refractum Aconitum subg. Aconitum Aconitum piepunense Aconitum legendrei Aconitum georgei Aconitum forrestii Aconitum chienningense Aconitum campylorrhynchum Aconitum bulleyanum Aconitum bulbilliferum Aconitum senanense Aconitum kitadakense 0.96 Aconitum nipponicum Aconitum hakusanense Aconitum jaluense Aconitum ciliare Aconitum yezoense Aconitum bartlettii Aconitum volubile Aconitum napiforme Aconitum kusnezoffii Aconitum carmichaelii Aconitum villosum 1.00 Aconitum variegatum 1.0 Aconitum napellus 0.99 Aconitum pentheri 1.00 1.0 Aconitum paskoi Aconitum baicalense 0.99 Aconitum yamazakii Aconitum apoiense Aconitum yuparense Aconitum sanyoense Aconitum iidemontanum Aconitum chiisanense Aconitum monanthum Aconitum austrokoreense Aconitum sungpanense 1.00 Aconitum racemulosum Aconitum anthora Aconitum polycarpum 0.99 Aconitum sinomontanum Aconitum monticola 1.0 Aconitum lycoctonum 0.99 Aconitum lasiostomum Aconitum shennongjiaense Aconitum septentrionale 1.0 Aconitum orientale Aconitum moldavicum 0.98 Aconitum umbrosum Aconitum gigas Aconitum subg. Lycoctonum Aconitum barbatum 0.98 Aconitum longecassidatum Aconitum finetianum 1.00 1.0 Aconitum alboviolaceum Aconitum scaposum Aconitum angustius 1.00 Aconitum chrysotrichum Aconitum crassiflorum 1.00 Delphinium staphisagria Delphinium pictum Delphinium subg. Staphisagria Delphinium requienii Megaleranthis saniculifolia Nigella damascena 0.5

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1.0 Aconitella aconiti 1.0 Aconitella thirkeana 1.0 Aconitella stenocarpa Aconitella hohenackeri Aconitella anthoroidea 1.0 Consolida pubescens 1.0 Consolida mauritanica 1.0 Consolida hellespontica Consolida glandulosa Aconitella barbata Consolida persica Consolida flava 1.0 Consolida stocksiana Consolida kabuliana 1.0 0.99 1.0 Consolida songorica Consolida 0.96 1.0 Consolida camptocarpa Consolida kandaharica Consolida aucheri 1.0 Consolida orientalis 1.00 Consolida hispanica 0.96 1.0 Consolida uechtritziana Consolida ajacis Consolida axilliflora Consolida regalis 1.0 Consolida lineolata Consolida armeniaca Consolida olopetala Delphinium tsarongense Delphinium orthocentrum Delphinium pachycentrum Delphinium sutchuenense Delphinium pyramidale Delphinium muscosum Delphinium kamaonense Delphinium grandiflorum Delphinium macrostachyum Delphinium kurdicum Delphinium freynii Delphinium fissum 0.96 Delphinium sylvaticum Delphinium emarginatum Delphinium pentagynum Delphinium nevadense Delphinium pylzowii Delphinium potaninii Delphinium delavayi Delphinium macrocentron Delphinium leroyi Delphinium dasycaulon Delphinium oreophilum Delphinium cashmerianum Delphinium wootonii Delphinium madrense Delphinium carolinianum 0.96 Delphinium oxysepalum Delphinium elatum Delphinium cuneatum Delphinium montanum Delphinium dubium Delphinium bulleyanum Delphinium tuberosum Delphinium szowitsianum Delphinium cyphoplectrum Delphinium aquilegifolium 0.99 Delphinium hesperium Delphinium hansenii Delphinium viridescens Delphinium glaucescens Delphinium nuttallianum 0.99 Delphinium lineapetalum 0.99 Delphinium depauperatum Delphinium menziesii Delphinium umbraculorum Delphinium recurvatum Delphinium decorum Delphinium gypsophilum Delphinium parryi Delphinium parishii Delphinium cardinale 0.98Delphinium hutchinsonae Delphinium californicum Delphinium multiplex Delphinium distichum 0.96 Delphinium subscandens Delphinium calcarequitis Delphinium viride Delphinium wislizeni Delphinium bicornutum 1.00 Delphinium sutherlandii Delphinium bicolor Delphinium xantholeucum Delphinium glareosum Delphinium nuttallii Delphinium glaucum Delphinium exaltatum Delphinium barbeyi Delphinium variegatum Delphinium scaposum Delphinium nudicaule Delphinium bakeri Delphinium subg. Delphinastrum Delphinium luteum Delphinium antoninum Delphinium trolliifolium 0.99 Delphinium geraniifolium Delphinium andesicola Delphinium ulignosum Delphinium stachydeum Delphinium polycladon Delphinium patens Delphinium inopinum Delphinium gracilentum Delphinium geyeri Delphinium andersonii Delphinium wendelboi Delphinium valens Delphinium tricorne Delphinium treleasei Delphinium tatsienense Delphinium tangkulaense Delphinium taliense Delphinium suave Delphinium spirocentrum Delphinium sparsiflorum Delphinium smithianum Delphinium semibarbatum Delphinium sapellonis Delphinium saniculifolium Delphinium ramosum Delphinium pycnocentrum Delphinium purpusii Delphinium pseudohamatum Delphinium omeiense Delphinium ochotense Delphinium novomexicanum Delphinium newtonianum Delphinium muliense Delphinium minjanense Delphinium middendorffii Delphinium maackianum Delphinium kohatense Delphinium kansuense Delphinium incisum Delphinium gyalanum Delphinium griseum Delphinium glaciale Delphinium flexuosum Delphinium elbursense Delphinium dolichostachyum Delphinium dolichocentroides Delphinium denudatum Delphinium crispulum Delphinium crassifolium Delphinium cheilanthum Delphinium ceratophorum Delphinium carduchorum Delphinium caeruleum Delphinium brachycentrum Delphinium bonvalotii 1.00 Delphinium biternatum Delphinium beesianum Delphinium batangense Delphinium alpestre Delphinium albocoeruleum Delphinium alabamicum Delphinium afghanicum Delphinium thibeticum Delphinium trichophorum Delphinium oxycentrum Delphinium forrestii continued Delphinium viscosum previous page 0.5

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99/1.00 Consolida axilliflora Consolida ajacis

100/1.00 Consolida persica Aconitella anthoroidea Consolida 100/0.99 Consolida regalis 100/ Consolida armeniaca Delphinium pseudohamatum 100/1.00 -/ Delphinium tatsienense Delphinium delavayi Delphinium Subg. Delphinastrum 91/1.00 100/1.00 Delphinium omeiense Delphinium anthriscifolium 76/0.99 100/1.00 Delphinium cossonianum Delphinium subg. Delphinium Delphinium virgatum Aconitum gymnandrum Aconitum subg. Gymnaconitum Aconitum pendulum Aconitum napellus Aconitum pentheri Aconitum ciliare Aconitum baicalense Aconitum subg. Aconitum 85/0.99 Aconitum ferox Aconitum delphinifolium 100/1.00 Aconitum racemulosum 100/ 1.00 Aconitum sinomontanum Aconitum septentrionale Aconitum monticola Aconitum barbatum Aconitum longecassidatum Aconitum subg. Lycoctonum 68/0.99 Aconitum alboviolaceum 76/0.99 Aconitum scaposum Aconitum angustius 100/1.00 Delphinium requienii Delphinium subg. Staphisagria Delphinium staphisagria Nigella damascena

0.01 Megaleranthis saniculifolia

Fig. S7. Phylogram of 34 accessions obtained from ML analysis of the cpDNA data. Numbers at the nodes are bootstrap percentages (> 70%) and Bayesian posterior probabilities (> 0.95), respectively. “-” indicates node not supported. Classification of Tamura (1995) is shown on the right.

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Consolida axilliflora Consolida ajacis Consolida regalis 100/1.00 Consolida persica Consolida 100/1.00 Aconitella anthoroidea

100/1.00 Consolida armeniaca Delphinium tatsienense 100/1.00 Delphinium pseudohamatum Delphinium omeiense Delphinium Subg. Delphinastrum Delphinium delavayi 89/0.99 100/1.00 Delphinium virgatum Delphinium cossonianum Delphinium subg. Delphinium Delphinium anthriscifolium Aconitum gymnandrum Aconitum subg. Gymnaconitum Aconitum pendulum Aconitum ferox 76/ Aconitum delphinifolium Aconitum ciliare 92/ Aconitum subg. Aconitum 0.99 Aconitum pentheri Aconitum napellus Aconitum baicalense Aconitum racemulosum

100/1.00 Aconitum sinomontanum 91/0.99 Aconitum monticola

100/1.00 Aconitum septentrionale Aconitum longecassidatum Aconitum subg. Lycoctonum Aconitum alboviolaceum

100/0.99 Aconitum barbatum Aconitum scaposum Aconitum angustius 100/1.00 Delphinium staphisagria Delphinium subg. Staphisagria Delphinium requienii Nigella damascena Megaleranthis saniculifolia 0.05

Fig. S8. Phylogram of 34 accessions obtained from BI analysis of the ITS data. Numbers at the nodes are bootstrap percentages (> 70%) and Bayesian posterior probabilities (> 0.95), respectively. Classification of Tamura (1995) is shown on the right.

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Appendix S1. Species and GenBank accession numbers for the datasets of two markers. “–” indicates missing data. Subgeneric classifications of Aconitum and Delphinium are based on Tamura (1995). Genus and/or subgenus (number of species studied/number of species in genus and/or subgenus); Species, voucher, locality, and GenBank accession: ITS and trnL-F. Ingroups: Aconitella Spach (6/~10): A. aconiti (L.) Soják, Buttler 20006 (M), Turkey: Konya, JF331874, JF331679; A. anthoroidea (Boiss.) Soják, Rechinger 42468 (M), Iran: Kurdistan, JF331875, JF331680; A. barbata (Bunge) Soják, Podlech 11320 (M), Afghanistan: Baghlan, JF331876, JF331681; A. hohenackeri (Boiss.) Soják, Nydegger 44486 (MSB), Turkey: Erzurum, JF331877, JF331682; A. stenocarpa (Hossain & P.H. Davis) Soják, Buttler 20012 (M), Turkey: Konya, JF331878, JF331685; A. thirkeana (Boiss.) Soják, Buttler & Erben 17987 (M), Turkey: Eskişehir, JF331879, JF331686; Aconitum L. (97/~300): subg. Lycocto- num (DC.) Peterm. (18/~40): A. alboviolaceum Kom., Yangqe0171 (PE), China: Hebei, JF975786, –; A. angustius W.T. Wang, Yangqe0185 (PE), China: Anhui, JF975796, –; A. barbatum var. puberulum Ledeb., Yangqe0206 (PE), China: Hebei, JF975804, –; A. chrysotrichum W.T. Wang, Y. Luo 358 (PE), China: Sichuan, AY164642, AY164649; A. crassiflorum Hand.-Mazz., Y. Luo 365 (PE), China: Sichuan, AY150230, AY150245; A. finetianum Hand.-Mazz., Q.E. Yang s.n. (PE), China: Jiangxi, AY164643, AY164650; A. gigas H. Lév. & Vaniot, 180011 (KANA), Japan: Hokkaido, AB004963, D86453; A. lasiostomum Rchb., 58429 (I), Rumania, AF216534, –; A. longecassidatum Nakai, Yangqe0198 (PE), China: Anhui, JF975809, –; A. lycoctonum L., M. Baltisberger & A. Widmer 13345 (ZT), Romania: Zarnasti, AF216537, Soininen & al. (2009), Norway, GQ244533; A. moldavicum Hacq., J. Sarbu 42108 (I), Rumania, AF216535, –; A. monticola Steinb., Yangqe0191 (PE), China: Hebei, JF975813, –; A. orientale Mill., B. Neuffer (OSBU), Russia: Teberda, AF216556, –; A. scaposum Franch., Y. Luo 327 (PE), China: Sichuan, AY150231, AY150246; A. septentrionale Koelle, Yarosh & Maslin ZZT5 (ZT), Russia: Ural, AF216552, Dietrich 5808 (M), Norway, JF331730; A. shennongjiaense Q. Gao & Q.E. Yang, Yangqe0220 (PE), China: Hubei, JF975819, –; A. sinomontanum Nakai, Y. Luo 333 (PE), China: Sichuan, AY150232, AY150247; A. umbrosum (Korsh.) Kom., Yangqe0163 (PE), China: Jilin, JF975825, –; subg. Aconitum (Stapf.) Rapaics (78/~250): A. anthora L., Utelli & Horat 14417 (ZT), France: Hautes-Pyrénées, AF216547, –; A. apoiense Nakai, 180075 (KANA), Japan: Hokkaido, AB004936, –; A. austro-koreense Koidz., Pak, Choi & Lee 4540, Korea: Taegu, AB020351, –; A. baicalense Turcz. ex Rapaics, 19027196 (TNS), Russia: Siberia, AB004941, –; A. baicalense Turcz. ex Rapaics, Zedtwitz 1936 (M), Russia: Transbaikal, –, JF331723; A. balfourii Stapf, Devarumath & al. (2004, unpub.), India, AJ717337, –; A. bartlettii Yamamoto, ITO Aco3, China: Taiwan, AB004946, –; A. brachypodum Diels, F.M. Zhang 00-007 (PE), China: Yunnan, AY189789, –; A. brunneum Hand.-Mazz., Yang 92025 (PE), China: Sichuan, AY571344, –; A. bulbilliferum Hand. -Mazz., Y. Luo & Y.S. Chen 548 (PE), China: Sichuan, AY571360, –; A. bulleyanum Diels, F.M. Zhang 00-016 (PE), China: Yunnan, AY189798, –; A. campylorrhynchum Hand. -Mazz., Y. Luo & Y. S. Chen 528 (PE), China: Sichuan, AY571359, –; A. cannabifolium Franch. ex Finet & Gagnep., 1391652 (PE), China: Anhui, AY189801, –; A. carmichaelii Debx., cult. Beijing Bot. G. (PE), AY571352, –; A. chienningense W.T. Wang, Y. Luo 353 (PE), China: Sichuan, AY164646, AY164653; A. chiisanense Nakai, Kita 951000 (TNS), Korea: Kyongsangnando, AB0049501, –; A. ciliare DC., Kita 951120, Japan: Kumamoto, AB004952, Herrmann 238 (M), China: Jilin, JF331724; A. columbianum Nutt., N. Tsipoura 22 (WS), U.S.A.: Idaho, AF258683, –; A. contortum Finet & Gagnep., F.M. Zhang 00-019 (PE), China: Yunnan, AY189788, –; A. decipiens Vorosch & Anfalov, Wakab & al. 9327102 (TNS) Russia: Siberia, AB004955, D86446; A. delavayi Franch., Zhang & Ge (2002, unpub.), no locality, AY189785, –; A. delphinifolium DC., Wells 1777 (WS), U.S.A.: Alaska, AF258681, Lang s.n. 28.07.1996 (M), no locality, JF331725; A. episcopale H. Lev., Y. Luo & Y.B. Luo 389 (PE), China: Sichuan, AY189794, –; A. fengii W.T. Wang, F.M. Zhang 00-003 (PE), China: Yunnan, AY189799, –; A. ferox Wall, Minaki & al. 9100909 (TI), Nepal: Mt. Shiwapuri, AB004961, J. Pcelt 29 (M), Nepal: Khumbu, JF331726; A. flavum Hand.-Mazz., Q.E. Yang 9501 (PE), China: Qinghai, AY571357, –; A. forrestii Stapf, F.M. Zhang 10005 (PE), China: Sichuan, AY164644, AY164651; A. franchetii Finet & Gagnepain, Y. Luo 390 (PE), China: Sichuan, AY150236, AY150251; A. georgei Comber, F.M. Zhang 00-012 (PE), China: Yunnan, AY189792, –; A. hakusanense Nakai, 180097 (KANA), Japan: Ishikawa, AB004964, –; A. hamatipetalum W. T. Wang, Y. Luo & Y.S. Chen 482 (PE), China: Yunnan, AY571355, –; A. hemsleyanum E. Pritz. ex Diels, Y. Luo 386 (PE), China: Sichuan, AY189793, AY150252; A. heterophyllum Wall, Balasubramani & al. (2011, unpulbished), India, JF440340, –; A. huiliense Hand.-Mazz., Y. Luo & Y.S. Chen 553 (PE), China: Sichuan, AY571362, –; A. iide-montanum Kadota, Kita & Ueda, 191948 (KANA), Japan: Yamagata, AB004970, D86456; A. jaluense Koma- rov, ITO Aco3 (no voucher), Korea: Kangwonda, AB004973, –; A. karakolicum Rapaics, D.Y. Tan A002 (PE), China: Xinjiang, AY571358, –; A. kitadakense Nakai, 191938 (KANA), Japan: Yamanashi, AB004977, D86463; A. kusnezoffii Rchb., Q.E. Yang s.n. (PE), China: Inner Mongolia, AY571346, –; A. legendrei Hand.-Mazz., Y. Luo & Y.S. Chen 448 (PE), China: Sichuan, AY571354, –; A. liangshanicum W.T. Wang, Y. Luo & Y.S. Chen 450 (PE), China: Sichuan, AY571364, –; A. liljestrandii Hand.-Mazz., Y. Luo & Y.B. Luo 344 (PE), China: Sichuan, AY164645, AY164652; A. monanthum Nakai, S. Ge 008 (PE), China: Jilin, AY189803, –; A. nagarum Stapf, Y.B. Luo 468 (PE), China: Yunnan, AY571347, Kadota 21473 (TNS), China: Yunnan, D86450; A. napellus L., Utelli & Soliva 14412 (ZT), Italy: Lago di Como, AF216544, –; A. napiforme Lev. & Vnt., 180103 (KANA), Japan: Nagano, AB004988, –; A. nemorum Popov, D.Y. Tan A001 (PE), China: Xinjiang, AY571361, –; A. nipponicum Nakai, 180103 (KANA), Japan: Nagano, AB004989, D86462; A. palmatum D.Don., Devarumath & al. (2004, unpub.), India, AJ717340, –; A. paskoi Vorosch., Wakabayashi & al. 9327216 (TNS), Russia: Siberia, AB004997, D86443; A. pendulum Busch, Y. Luo & Y.B. Luo 411 (PE), China: Sichuan, AY150235, D. Boufford & al. 40471 (MSB), China: Sichuan, JF331728; A. pentheri Hayek, D. Podlech & W. Lippert 26275 (M), Serbia, JF331905, JF331729; A. piepunense Hand.-Mazz., F.M. Zhang 00-001 (PE), China: Yunnan, AY189800, –; A. pilopetalum W.T. Wang & L.Q. Li, Y. Luo & Y.B. Luo 364 (PE), China: Sichuan, AY571343, –; A. polycarpum Chang ex W.T. Wang, Y. Luo & Y.S. Chen 494 (PE), China: Yunnan, AY571356, –; A. pulchellum Hand.-Mazz., Y. Luo & Y.B. Luo 362 (PE), China: Sichuan, AY164647, AY164654; A. racemulosum Franch., Y. Luo & Y.B. Luo 360 (PE), China: Sichuan, AY150233, Wang Wei 081 (PE), China: Chongqing, FJ626533; A. refractum (Finet & Gagnep.) Hand.-Mazz., Y. Luo & Y.B. Luo 406 (PE), China: Sichuan, AY571349, –; A. sanyoense Nakai, 180095 (KANA), Japan: Kyoto, AB005002, D86263; A. senanense Nakai, 191943 (KANA), Japan: Yamanashi, AB005006, D86461; A. sessiliflorum (Finet & Gagnep.) Hand.-Mazz., Y. Luo 396 (PE), China: Sichuan, AY164648, AY164655; A. spiripetalum Hand.-Mazz., Y. Luo & Y.B. Luo 393 (PE), China: Sichuan, AY571345, –; A. stapfianum Hand.-Mazz., Zhang & Ge (2002, unpublised), no locality, AY189786, –; A. sungpanense Hand.-Mazz., H.Z. Kong k1001 (PE), China: Shaanxi, AY189795, –; A. tanguticum (Maxim.) Stapf, Y. Luo & Y.B. Luo 309 (PE), China: Sichuan, AY150234, AY150249; A. tatsienense Finet & Gagnep., Y. Luo & Y.B. Luo 384 (PE), China: Sichuan, AY571348, –; A. tongolense Ulbr., Y. Luo & Y.B. Luo 375 (PE), China: Sichuan, AY571350, –; A. transsectum Diels, F.M. Zhang 00-011 (PE), China: Yunnan, AY189791, –; A. tsaii W.T. Wang, F.M. Zhang 00-014 (PE), China: Yunnan, AY189784, –; A. tuguancunense Q.E. Yang, Zhang & Ge (2002, unpub.), China, AY189782, –; A. variegatum L., A.B. Utelli 14428 (ZT), Switzerland: Bergell, AF216549, –; A. villosum Rchb., 9237144 (TNS), Russia: Siberia, AB005010, D86442; A. vilmorinianum Kom, Zhang & Ge (2002, unpub.), China, AY189787, –; A. violaceum Jacquem. ex Stapf, Devarumath & al. (2004, unpub.), India, AJ717338, Valentini & al. (2009), no locality, EU326032; A. volubile Muhl., S. Ge 004 (PE), China: Jilin, AY189802, –; A. yamazakii Tamura & Namba, 180040 (KANA), Japan: Hokkaido, AB005012, D86260; A. yezoense Nakai, 180045 (KANA), Japan: Hakkaido, AB005018, –; A. yuparense Takeda, 180022 (KANA), Japan: Hakkaido, AB005019, D86262; A. zeravschanicum Steinb., Kita 9532187 (TNS), Kirghiz, AB005022, –; subg. Gymnaconitum (Stapf.) Rapaics (1/1): A. gymnandrum Maxim., Y. Luo 326 (PE), China: Sichuan, AY150238, AY164651; Consolida (DC.) S.F. Gray (21/~40): C. ajacis (L.) Schur, Merxmüller 8524 (M), Germany: Bavaria, JF331880, JF331687; C. armeniaca (Stapf ex Huth) Schröd., Nydegger 16993 (MSB), Turkey, JF331883, JF331690; C. aucheri (Boiss.) Iranshahr, Podlech & Jarmal 28879 (MSB), Afghanistan: Kandahar, JF331884, JF331691; C. axilliflora (DC.) Schrödinger, Brachfeld & Graben 42414 (MSB), Turkey: Anatolia, JF331885, JF331692; C. camptocarpa (Fisch. & C.A. Mey.) Nevski, Rusanovitsch & Kramarenko 14029 (M), Kazakhstan: Taldy-Kurgan, JF331886, JF331694; C. flava (DC.) Schrödinger ex Hand.-Mazz., Rechinger 148 (M), Iraq: Haswa desert, JF331887, JF331695; C. glandulosa (Boiss. & A. Huet) Bornm., Nydegger 16741 (M), Turkey: Anatolia, JF331888, JF331696; C. hellespontica (Boiss.) Chater, Nydegger 44206 (MSB), Turkey: Konya, JF331889, JF331697; C. hispanica (Willk. ex Costa) Greuter & Burdet, Angerer s.n. 25.7.1984 (M), Germany: Bavaria, JF331890, JF331698; C. kabuliana (Akhtar) Iranshahr, Volk 2630 (M), Afghanistan: Kabul, JF331891, JF331700; C. kandaharica Iranshahr, Podlech 22009 (MSB), Afghanistan: Farah, JF331892, JF331701; C. lineolata Hub.-Mor. & C. Simon, Nydegger 13313 (MSB), Turkey, JF331893, JF331703; C. mauritanica (Coss.) Munz, Podlech 47566 (MSB), Morocco: Er-Rachidia, JF331894, JF331704; C. olopetala (Boiss.) Hayek, Nydegger 43795 (MSB), Turkey: Erzincan, JF331895, JF331706; C. orientalis (J. Gay) Schrödinger, Parishani 14382 (M), Iran:

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Appendix S1. Continued. Isfahan, JF331896, JF331707; C. persica (Boiss.) Schroedinger, Rechinger 42445 (M), Iran: Kurdistan, JF331897, JF331708; C. pubescens (DC.) Soó, Lambinon 79E568 (MSB), Spain: Zaragoza, JF331899, JF331710; C. regalis Gray, Hertel 6560 (M), Austria: Parndorf plain, JF331900, JF331714; C. songorica Nevski, Karelin & Kiriloff 1165 (MSB), Kazakhstan, JF331902, JF331719; C. stocksiana (Boiss.) Nevski, Volk 1585 (M), Afghanistan: Kabul, JF331903, JF331720; C. uechtritziana (Huth) Soó, Rechinger 21108 (M), Greece: Epirus, JF331904, JF331722; Delphinium L. (161/~320): subg. Delphinastrum (DC.) Peterm. (143/~300): D. afghanicum Rech. f., Grözbach 1 (MSB), Afghanistan: Takhar, –, JN573529; D. alabamicum Kral, Bryson 3150 (GA), U.S.A.: Alabama, AF258756, –; D. albocoeruleum Maxim., Trippner 191 (M), China: Kansu, –, JN573530; D. alpestre Rydberg, Hogan 2923 (COLO), U.S.A.: Colorado, AF258733, –; D. andersonii A. Gray, Richter 7 (WS), U.S.A.: Oregon, AF258773, AF258647; D. andesicola Ewan, Fishbein 618 (ARIZ), Mexico: Sonora, AF258725, –; D. antoninum Eastw., Jokerst 2433 (CHSC), U.S.A.: California, AF258691, –; D. aquilegifolium (Boiss.) Bornm., Manutsheri 1066 (MSB), Iran: Mazanderan, –, JN573531; D. bakeri Ewan, Snow 1230 (RPBG), U.S.A.: California, AF258697, AF258652; D. barbeyi Huth, Siplivisky & Beck 4314 (WS), U.S.A.: Colorado, AF258709, AF258639; D. batangense Finet & Gagnep., Yangqe0036 (PE), China: Yunnan, JF976224, –; D. beesianum W.W. Sm., Farrer & Purdom 237 (M) China: Kansu, –, JN573532; D. bicolor Nutt., P. Soltis 2383 (WS), U.S.A.: Idaho, AF258711, AF258653; D. bicornutum Hemsl., Mendoza 1905 (ARIZ), Mexico: Chihuahua, AF258701, –; D. biternatum Huth, D. Podlech 15909 (M), Afghanistan: Baghlan, –, JN573533; D. bonvalotii Franch., Wang Wei 030 (PE), China: Guizhou, –, FJ626542; D. brachycentrum Ledeb., Voron & Derviz-Sokolova s.n. 1967 (M), Russia: Magadan, –, JN573534; D. bulleyanum Forrest ex Diels, cult. Munich Bot. G., China: Yunnan, –, JN573535; D. caeruleum Jacquem., J. Pcelt s.n. 19.09.1962 (M), Nepal: Khumbu, –, JN573536; D. calcarequitis Standl., Ferguson (ARIZ), Mexico: Chihuahua, AF258719, –; D. californicum Torr., San Mateo s.n. (RPBG), U.S.A.: California, AF258713, AF258640; D. cardinale Hook., Mort 1374 (no voucher), U.S.A.: California, AF258740, AF258648; D. carduchorum Chowdhuri & P.H. Davis, Vogel s.n. 16.08.1968 (M), Turkey: Hakkari, –, JN573537; D. carolinianum subsp. vimineum (D. Don) M.J. Warnock, Thomas 154175 (WS), U.S.A.: Louisiana, AF258720, –; D. cashmerianum Royle, Anders 8118 (MSB), Afghanistan: Badakhstan, –, JN573538; D. ceratophorum Franch., Yangqe0023 (PE), China: Yunnan, JF976226, –; D. cheilanthum Fisch. ex DC., Leuchtenb s.n (M), Russia: Siberia, –, JN573539; D. crassifolium Schrad. ex Spreng., Boyko & Starch. s.n. (M), Russia: Amur, –, JN573540; D. crispulum Rupr., K.H. Rechinger 57118 (M), Azerbaijan: Mt. Sahand, –, JN573541; D. cuneatum Steven ex DC., Litvinov 7219 (MSB), Russia: Samara, –, JN573542; D. cyphoplectrum Boiss., S. Zarre & Zarrei 1042 (MSB), Iran: Bushehr, –, JN573543; D. dasycaulon Fresen, Mbala 592 (M), Zambia, –, JN573544; D. decorum Fisch. & C.A. Mey., J. Koontz 63 (WS), U.S.A.: California, AF258744, AF258631; D. delavayi Franch., McBeath & al. CLD0895 (UCBG), China: Yulong Shan, AF258705, AF258659; D. denudatum Wall., D. Podlech 31929 (MSB), Afghanistan: Ghazni, –, JN573546; D. depauperatum Nutt., Richter 59 (WS), U.S.A.: Oregon, AF258689, AF258656; D. distichum Geyer ex Hooker, Fishbein 3460 (WS), U.S.A.: Washington, AF258774, –; D. dolichocentroides W.T. Wang, Yangqe0025 (PE), China: Yunnan, JF976231, –; D. dolichostachyum Chowdhuri & P.H. Davis, Thoma s.n. (M), Turkey: Munzur-Gebirge, JN573516, JN573547; D. dubium (Rouy & Fouc.) Pawl., H. Merxmüller & Wiedmann 285/64 (M), Italy: Cuneo, –, JN573548; D. elatum L., Skvortsov & al. 10597 (M), Russia: Ural, JN5735179, JN57354; D. elbursense Rech. f., Manutshehri 983 (MSB), Iran: Gilan, –, JN573550; D. emarginatum C. Presl, H. Ross 303 (M), Italy: Blermo, –, JN573551; D. exaltatum Aiton, Bright 18948 (WS), U.S.A.: Pennsylvania, –, AF258651; D. fissum Waldst. & Kit., K.H. Rechinger 21506 (M), Greece: Epirus, –, JN573552; D. flexuosum M. Bieb., Gagnide & al. 887 (MO), Georgia: Khevi, –, JN573553; D. forrestii Diels, Yangqe0020 (PE), China: Yunnan, JF976232, –; D. freynii Huth, G. Fayvush 1631 (M), Armenia: Vayotsdzor, –, JN573554; D. geraniifolium Rydberg, Wallace s.n. (ARIZ), U.S.A.: Arizoma, AF258765, –; D. geyeri Greene, Nelson & Nelson 6702 (WS), U.S.A.: Wyoming, AF258762, –; D. glaciale Hook. f. & Thomson, Miyamoto & al. 9592453 (MO), Nepal: Sagarmatha, –, JN573555; D. glareosum Greene, Grable 5772 (WS), U.S.A.: Washington, AF258764, –; D. glaucescens Rydberg, Hitckcock & Sutherland 2351 (WS), U.S.A., Idaho, AF258754, AF258645; D. glaucum S. Watson, Cheatham s.n. (WS), U.S.A.: Oregon, AF258739, –; D. gracilentum Greene, Warnock s.n. (WS), U.S.A.: California, AF258763, –; D. grandiflorum L., Erskine & al. SICH205 (UCBG), China: Sichuan, AF258761, AF258630; D. griseum Gilli, Volk 1873 (M), Afghanistan: Kabul, –, JN573556; D. gyalanum C. Marquand & Airy Shaw, cult. Munich Bot. G., China: Tibet, –, JN573557; D. gypsophilum Ewan, Koontz 36 (WS), U.S.A.: Washington, AF258721, AF258633; D. hansenii Greene, Warnock s.n. (WS), U.S.A.: California, AF258760, –; D. hesperium A. Gray, Raiche 20005 (UCBG), U.S.A.: California, AF258772, AF258634; D. hutchinsonae Ewan, 94.123 (RPBG), U.S.A.: California, AF258750, AF258641; D. incisum Wall., J. Pcelt s.n. (M), Nepal: Okhaldunga, –, JN573558; D. inopinum (Jepson) H.F. Lewis & Epling, Grable 11320 (WS), U.S.A.: California, AF258749, –; D. kamaonense Huth, Yangqe0048 (PE), China: Sichuan, JF976235, Wuendish 303 (MSB), Nepal: Langtang, JN573559; D. kansuense W.T. Wang, Q.E. Yang & Q. Yuan 386 (IBSC), China: Fin’an, –, JN573560; D. kohatense (Brühl) Munz, Anders 3567 (MSB), Afghan- istan: Paktia, –, JN573561; D. kurdicum Boiss. & Hohen., K.H. Rechinger 11017 (M), Iraq: Erbil, –, JN573562; D. leroyi Franch. ex Huth, Richards 25743 (M), Tanzania: Arusha, –, JN573563; D. lineapetalum Ewan, Fishbein 3211 (WS), U.S.A.: Washington, AF258771, –; D. luteum A. Heller, Guggolz 1542 (WS), U.S.A.: California, AF258777, –; D. maackianum Regel, Zedtwitz s.n., (M), Russia, –, JN573565; D. macrocentron Oliv., W. Rauh 410 (M), Kenya, –, JN573566; D. macrostachyum Boiss. ex Huth, K.H. Rechinger 43020 (M), Iran: Kurdistan, –, JN573568; D. madrense S. Watson, Warnock 2252 (LL), U.S.A.: Texas, AF258712, –; D. menziesii de Candolle, Richter 5 (WS), U.S.A.: Oregon, AF258755, –; D. middendorffii Trautv., Korol. & Rjab. 6025 (M), Russia: Magadan, –, JN573569; D. minjanense Rech. f., Breckle 2998 (MSB), Afghanistan: Kunar, –, JN573570; D. montanum DC., Kalheber 85-2044 (M), France: Pyrénées Orientales, JN573518, N573571; D. muliense W.T. Wang, Yangqe0025 (PE), China: Yunnan, JF976237, –; D. multiplex (Ewan) C.L. Hitchc., Hitchcock & Muhlick s.n. (WS), U.S.A.: Washington, AF258714, AF258654; D. muscosum Exell & Hillc., A. Kress s.n. (M), –, JN573519, JN573572; D. nevadense Kunze, Bertel 11109 (M), Spain: Granada, –, JN573574; D. newtonianum D.M. Moore, Rettig 1145 (GA), U.S.A.: Arkansas, AF258742, –; D. novomexicanum Wooton, Worthington 4771 (ARIZ), U.S.A.: New Mexico, AF258718, –; D. nudicaule Torr. & A. Gray, J. Koontz 96-20 (WS), U.S.A.: California, AF258728, AF258638; D. nuttallianum Pritz., Strickler 91/2-6 (WS), U.S.A.: Washington, AF258688, AF258646; D. nuttallii A. Gray, K. Karoly s.n. (OR), U.S.A.: Oregon, AF258746, –; D. ochotense Nevski, Soininen & al. (2009), Norway, –, GQ244830; D. omeiense W.T. Wang, Yangqe0043 (PE), China: Sichuan, JF976239, –; D. oreophilum Huth, Vassiljeva s.n. (M), Uzbekistan, –, JN573576; D. orthocentrum Franch., Y. Luo 323 (PE), China: Sichuan, AY150242, AY150257; D. oxycentrum W.T. Wang, H.Z. Kong 98-329 (PE), China: Yunnan, AY150240, AY150255; D. oxysepalum Pax & Borbás, Angerer s.n. (M), Slovakia, JN573522, JN573577; D. pachycentrum Hemsl. ex Brühl, Y. Luo 311 (PE), China: Sichuan, AY150243, AY150258; D. parishii A. Gray, Richter 53 (WS), U.S.A.: California, AF258716, AF258635; D. parryi A. Gray, Edwards s.n. (RPB), U.S.A.: California, AF258694, AF258636; D. patens Benth., Richter 11 (WS), U.S.A.: California, AF258734, AF258658; D. pentagynum Lam., Lewalle 10967 (MSB), Morocco: Rabat, JN573523, JN573578; D. polycladon Eastw., R.C. Bacigalupi 6508 (WS), U.S.A.: California, AF258743, AF258642; D. potaninii Huth, Yangqe0005 (PE), China: Sichuan, JF976245, –; D. potaninii Huth, Q.E. Yang & Q. Yuan 805 (IBSC), China: Zhuxi, –, JN573579; D. pseudohamatum W.T. Wang, Yangqe0027 (PE), China: Sichuan, JF976246, –; D. purpusii Brandegee, Shevock 12032 (RSA), U.S.A.: California, AF258747, –; D. pycnocentrum Franch., Yangqe0055 (PE), China: Yunnan, JF976249, –; D. pylzowii Maxim., Farrer & Purdom 253 (M), China: Kansu, –, JN573580; D. pyramidale Royle, A. Kress s.n. (M), –, JN573524, JN573581; D. ramosum Rydb., Hogan 3563 (COLO), U.S.A.: Colorado, AF258687, –; D. recurvatum Greene, Brown s.n. (WS), U.S.A.: California, AF258775, AF258632; D. saniculifolium Boiss., K.H. Rechinger 3838 (M), Iran: Kerman, –, JN573583; D. sapellonis Cockerell, Lowery 1877 (WS), U.S.A.: New Mexico, AF258708, –; D. scaposum Greene, Joyce 136 (WS), U.S.A.: Arizona, AF258732, AF258649; D. semibarbatum Bien. ex Boiss., Belianina & al. 9174 (MSB), Russia: Kuhitang Mtns, –, JN573584; D. smithianum Hand.-Mazz., Yangqe0050 (PE), China: Sichuan, JF976250, –; D. sparsiflorum Maxim., Trippner s.n. (M), China: Kansu, JN573525, JN573585; D. spirocentrum Hand.- Mazz., Yangqe0022 (PE), China: Yunnan, JF976253, –; D. stachydeum (A. Gray) Tidestr., Richter 61 (WS), U.S.A.: Oregon, AF258684, AF258643; D. suave Huth, Peer s.n. (MSB), Pakistan: Chitral, –, JN573586; D. subscandens Ewan, Correll & Gentry 23019 (LL), Mexico: Chihuahua, AF258710, –; D. sutchuenense Franch., D. Boufford & al. 39837 (MSB), China: Sichuan, JN573526, JN573587; D. sutherlandii M.J. Warnock, Bjork 1187 (WS), U.S.A.: Idaho, AF258693, –; D. sylvaticum Turcz., Lager s.n. (G), Algeria: Kabylia, Alger, –, JN573588; D. szowitsianum Boiss., K.H. Rechinger 43500 (M), Azerbaijan: Khalkal, –, JN573589; D. taliense Franch., Yangqe0009 (PE), China: Yunnan, JF976257, –; D. tangkulaense W.T. Wang, Q.E. Yang & Q. Yuan 409 (IBSC), China: Madoi, –, JN573590; D. tatsienense Franch., Yangqe0033 (PE), China: Sichuan, JF976259, A. Kress s.n. (M), China, JN573591; D. thibeticum Finet & Gagnep., Yangqe0014 (PE), China: Yunnan, JF976263, Rock 17338 (M), China: Yunnan, JN573592; D. treleasei Bush ex K.C. Davis, Brooks 14186 (GA), U.S.A.:

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Appendix S1. Continued. Missouri, AF258768, –; D. trichophorum Franch., Yangqe0016 (PE), China: Sichuan, JF976266, D. Boufford & al. 39746 (MSB), China: Sichuan, JN573593; D. tricorne Michx., Vincent 7734 (WS), U.S.A.: Ohio, AF258770, –; D. trolliifolium A. Gray, Edwards s.n. (RPBG), U.S.A.: California, AF258686, AF258650; D. tsarongense Hand.-Mazz., Handel-Mazzetti 7934 (M), China: Tibet, –, JN573594; D. tuberosum Aucher ex Boiss., K.H. Rechinger 47131 (M), China: Sich- uan, –, JN573595; D. ulignosum Curran, Grable 11242 (WS), U.S.A.: California, AF258766, –; D. umbraculorum F.H. Lewis & Epling, Warnock s.n. (WS), U.S.A.: California, AF258752, –; D. valens Standl., Nesom 7135 (LL), Mexico: Nuevo Leon, AF258703, –; D. variegatum Torr. & A. Gray, Richter 14 (WS), U.S.A.: California, AF258724, –; D. viride S. Watson, Nesom 4969 (TEX/L), Mexico: Chihuahua, AF258685, –; D. viridescens Leiberg, Naas & Naas 4019 (WS), U.S.A.: California, AF258736, AF258644; D. viscosum Hook. f. & Thomson, J. Pcelt s.n. (M), Nepal: Okhaldunga, JN573528, JN573597; D. wendelboi Iranshahr, Anders 6143 (MSB), Afghanistan: Baghlan, –, JN573598; D. wislizeni Engelm., Trauba s.n. (ARIZ), Mexico: Sonora, AF258735, –; D. wootonii Rydb., Fishbein 376a (ARIZ), U.S.A.: Arizona, AF258690, –; D. xantholeucum Piper, Hitchcock 17293 (WS), U.S.A.: Washington, AF258695, –; subg. Staphisagria (DC.) Peterm. (3/3): D. pictum Willd., Rumsey 15012 (M), Balearic Islands: Majorca, –, JF331741; D. requienii DC., Gavelle s.n. (M), France: Porquerolles Island, JF331742, JN573582; D. staphisagria L., Vitek 02-205 (M), Greece: Crete, JF332022, JF331743; subg. Delphinium W.T. Wang (15/~18): D. anthriscifolium Hance, D. Podlech 55468 (MSB), China: Shanxi, JF331919, JF331731; D. balansae Boiss. & Reut., Sammet & Illitz s.n., 1991 (MSB), Mo- rocco: Marrakech, JF331931, JF331732; D. balcanicum Pawl., Rösler 6570 (M), Serbia, JF331945, JF331733; D. cossonianum Batt., D. Podlech 46655 (MSB), Morocco: Meknes, JF331955, JF331734; D. favargeri C. Blanché, Molero & Simon Pall., Jury & Ait Lafkih 19781 (M), Morocco: Middle Atlas, JF331965, JF331735; D. gracile DC., Lambinon 79/E/534 (MSB), Spain: Teruel, JF331977, JF331736; D. halteratum Sibth. & Sm., Dunkel MTB3647.4 (M), Italy: L’Aquila, JF331982, JF331737; D. hirschfeldianum Heldr. & Holzm. ex Boiss., No author 1604 (M), Greece: Mykonos, JF331988, JF331738; D. macropetalum DC., D. Podlech 55412 (MSB), Morocco, JF331996, JF331739; D. nanum DC., Zubizaretta 42606 (M), Spain: Marbella, JN573520, JN573573; D. obcordatum DC., Nowotny 61.12.18 (M), Spain: Torremolinos, JN573521, JN573575; D. peregrinum L., Franzén & al. 870 (M), Macedonia, JF332001, JF331740; D. venulosum Boiss., Nydegger 15452 (MSB), Turkey, JF332024, JF331744; D. verdunense Balb., Krach & Koepff 3814 (MSB), Morocco: Meknes, JN573527, JN573596; D. virgatum Poir., S. Zarre 53 (MSB), Turkey: Kayseri, JF332030, JF331745. — Outgroups: Megaleranthis saniculifolia Ohwi, K. Heo 550 (KWNU), Korea, AY515399, Won JY 950520 (PE), HQ440188; Nigella damascena L., cult. Mainz Bot. G. No. 041773 (MJG), EU699446, Luo & al. (2002, unpub.), AY150260.

Appendix 2. Species, their sources, localities, and GenBank accession numbers for the datasets of six markers (ITS, trnL-F, trnH-psbA, trnK-matK, trnS- trnG, rbcL). “–” indicates missing data. Subgeneric classifications of Aconitum and Delphinium are based on Tamura (1995). Ingroups: Aconitella Spach: A. anthoroidea (Boiss.) Soják, Rechinger 42468 (M), Iran: Kurdistan, JF331875, JF331680, –, JF331747, JF331811, JF331666; Aconitum L.: subg. Lycoctonum (DC.) Peterm.: A. alboviolaceum Kom., Yangqe0171 (PE), China: Hebei, JF975786, –, JN043735, JF953007, –, JF940640; A. angustius W.T. Wang, Yangqe0185 (PE), China: Anhui, JF975796, –, JN043745, JF953017, –, JF940650; A. barbatum var. puberulum Ledeb., Yangqe0206 (PE), China: Hebei, JF975804, –, JN043753, JF953025, –, JF940657; A. longecassidatum Nakai, Yangqe0198 (PE), China: Anhui, JF975809, –, JN043759, JF953031, –, JF940663; A. monticola Steinb., Yangqe0191 (PE), China: Hebei, JF975813, –, JN043764, JF953035, –, JF940668; A. scaposum Franch., Y. Luo 327 (PE), China: Sichuan, AY150231, AY150246, Yangqe0168 (PE), China: Gansu, JN043769, JF953039, –, JF940674; A. septentrionale Koelle, Yarosh & Maslin ZZT5 (ZT), Russia: Ural, AF216552, Dietrich 5808 (M), Norway, JF331730, Yarosh & Maslin ZZT5 (ZT), Russia: Ural, AF216575, Dietrich 5808 (M), Norway, JF331795, JF331859, JF331678; A. sinomontanum Nakai, Y. Luo 333 (PE), China: Sichuan, AY150232, AY150247, Yangqe0166 (PE), China: Hebei, JN043775, JF953043, –, JF940678; subg. Aconitum (Stapf.) Rapaics: A. baicalense Turcz. ex Rapaics, 19027196 (TNS), Russia: Siberia, AB004941, Graf Zedtwitz 1936 (M), Russia: Transbaikal, JF331723, –, JF331788, JF331852, –; A. ciliare DC., Kita 951120, Japan: Kumamoto, AB004952, Herrmann 238 (M), China: Jilin, JF331724, –, JF331789, JF331853, –; A. delphinifolium DC., Wells 1777 (WS), U.S.A.: Alaska, AF258681, Lang s.n. 28.07.1996 (M), no locality, JF331725, –, JF331790, JF331854, –; A. ferox Wall, Minaki & al. 9100909 (TI), Nepal: Mt. Shiwapuri, AB004961, J. Pcelt 29 (M), Nepal: Khumbu, JF331726, –, JF331791, JF331855, –; A. napellus L., Utelli & Soliva 14412 (ZT), Italy: Lago di Como, AF216544, –, AF216567, zpl 01675 (MIB), Italy: Lombardy, FN668831, –, S. Hoot 926 (UWM), no locality, EU053898; A. pendulum Busch, Y. Luo & Y.B. Luo 411 (PE), China: Sichuan, AY150235, D. Boufford & al. 40471 (MSB), China: Sichuan, JF331728, –, JF331793, JF331857, –; A. pentheri Hayek, D. Podlech & W. Lippert 26275 (M), Serbia, JF331905, JF331729, –, JF331794, JF331858, –; A. racemulosum Franch., Y. Luo & Y.B. Luo 360 (PE), China: Sichuan, AY150233, Wang Wei 081 (PE), China: Chongqing, FJ626533, –, FJ626484, –, AY954488; subg. Gymnaconitum (Stapf.) Rapaics: A. gymnandrum Maxim., Y. Luo 326 (PE), China: Sichuan, AY150238, JF331727, LIDZ-1287 (KUN), no locality, GQ337734, Dickoré 9111 (MSB), China: Xizang, JF331792, JF331856, JF331677; Consolida (DC.) S.F. Gray: C. ajacis (L.) Schur, Merxmüller 8524 (M), Ger- many: Bavaria, JF331880, JF331687, M. Baltisberger & D. Frey (ZT), cult. in Zurich, AF216578, Merxmüller 8524 (M), Germany: Bavaria, JF331754, JF331818, JF331667; C. armeniaca (Stapf ex Huth) Schröd., Nydegger 16993 (MSB), Turkey, JF331883, JF331690, –, JF331757, JF331821, JF331672; C. axilliflora (DC.) Schrödinger, Brachfeld & Graben 42414 (MSB), Turkey: Anatolia, JF331885, JF331692, –, JF331759, JF331823, JF331669; C. persica (Boiss.) Schroedinger, Rechinger 42445 (M), Iran: Kurdistan, JF331897, JF331708, –, JF331773, JF331837, JF331673; C. regalis Gray, Hertel 6560 (M), Austria: Parndorf plain, JF331900, JF331714, –, JF331778, JF331842, JF331671; Delphinium L.: subg. Delphinastrum (DC.) Peterm.: D. delavayi Franch., McBeath & al. CLD0895 (UCBG), China: Yulong Shan, AF258705, AF258659, Yangqe0002 (PE), China: Yunnan, JN044403, –, –, JF941393; D. omeiense W.T. Wang, Yangqe0043 (PE), China: Sichuan, JF976239, –, JN044414, JF953657, –, JF941405; D. pseudohamatum W.T. Wang, Yangqe0027 (PE), China: Sichuan, JF976246, –, JN044420, JF953659, –, JF941413; D. tatsienense Franch., Yangqe0033 (PE), China: Sichuan, JF976259, JN573591, JN044434, JF953662, –, JF941426; subg. Staphisagria (DC.) Peterm.: D. requienii DC., Gavelle s.n. (M), France: Porquerolles Island, JF331742, JN573582, –, JF331806, JF331870, JF332021; D. staphisagria L., Vitek 02-205 (M), Greece: Crete, JF332022, JF331743, –, JF331807, JF331871, –; subg. Delphinium W.T. Wang: D. anthriscifolium Hance, D. Podlech 55468 (MSB), China: Shanxi, JF331919, JF331731, –, JF331796, JF331860, JF331674; D. cossonianum Batt., D. Podlech 46655 (MSB), Morocco: Meknes, JF331955, JF331734, –, JF331799, JF331863, JF331675; D. virgatum Poir., S. Zarre 53 (MSB), Turkey: Kayseri, JF332030, JF331745, –, JF331809, JF331873, –. — Outgroups: Megaleranthis saniculifolia Ohwi, K. Heo 550 (KWNU), Korea, AY515399, Won JY 950520 (PE), HQ440188, K.J. Kim s.n. (BCI), NC_012615, NC_012615, NC_012615, NC_012615; Nigella damascena L., cult. Mainz Bot. G. No. 041773 (MJG), EU699446, Luo & al. (2002, unpub.), AY150260, –, Wang Wei Seed1 (PE), China: Yunnan, FJ626504, –, FJ626586.

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