AbstractScorzonera comprises 180–190 species and belongs to the subtribe Scorzonerinae. Its circumscription has long been the subject of debate and available molecular phylogenetic analyses affirmed the polyphyly of Scorzonera in its wide sense. We provide a re-evaluation of Scorzonera and other related genera, based on carpological  and extended molecular phylogenetic analyses. We present, for the first time, a comprehensive sampling, including Scorzonera in its widest sense and all other genera recognised in the Scorzonerinae. We conducted phylogenetic analyses using Maximum Parsimony, Maximum Likelihood and Bayesian analyses, based on sequences of the nuclear ribosomal ITS and of two plastid markers  and Maximum Parsimony for reconstructing the carpological character states at ancestral nodes. Achene characters, especially related to pericarp anatomy, such as general topography of the tissue types, disposition of the mechanical tissue and direction of its fibres, presence or absence of air cavities, provide, in certain cases, support for the phylogenetic lineages revealed. Confirming the polyphyly of Scorzonera, we propose a revised classification of the subtribe, accepting the genera Scorzonera , Gelasia, Lipschitziagen. nov. (for the Scorzoneradivaricata clade), Pseudopodospermum, Pterachaenia , Ramaliellagen. nov. (for the S.polyclada clade) and Takhtajaniantha. A key to the revised genera and a characterisation of the genera and major clades are provided. Scorzonera L. with some 180–190 species is the largest and name-giving genus of the subtribe Scorzonerinae of the Cichorieae  of bristles with long soft and often intertwining fimbriae , Gelasia  are separated into lineages with different basic chromosome numbers  to investigate the variation in achene and pappus morphology and anatomy across the entire subtribe, to define carpological characters and states and to optimise them on to the new molecular phylogenetic tree, in order to assess the correspondence of carpological features with the principal clades and to gain insights into the evolution of carpological characters in the subtribe; (3) to characterise clades using carpological and, where available, further morphological and cytological characters, to review and, where sufficient evidence is provided, revise the current generic classification of the subtribe.The aims of the present study are (1) to provide a molecular phylogenetic analysis, using both nuclear and plastid DNA markers and a sampling that spans the entire subtribe, as well as the various groups of Scorzonera in its widest sense, as well as representatives of all other genera of the subtribe Scorzonerinae  gene; (3) a 750 bp fragment of the plastid DNA maturase K gene (matK).Sequences of three markers were used for the molecular phylogenetic analyses: (1) the nrITS region ; (2) a ~570 bp fragment of the plastid DNA ribulose 1,5-bisphosphate carboxylase/oxygenase large subunit . A two-step PCR method was used for library preparation: first-stage PCR using fusion primers containing the primer sequences from rbcL marker was sequenced on the 454 platform . The three-step PCR method was used for library preparation. All PCR stages were conducted using Phusion high fidelity DNA-polymerase . The first-stage PCR was performed using a reaction mixture of a total volume of 20 µl: 4 μl 5× buffer for Phusion high fidelity DNA-polymerase, 250 µM dNTP , 0.2 μl Phusion high fidelity DNA-polymerase and 0.8 pM each primers , 0.2 μl of Phusion high fidelity DNA-polymerase, 0.3 pM of each primer, 4 μl of amplification products of the previous stage. PCR conditions: 98 °C – 1 min; 18 cycles each for 98 °C – 5 s, 55 °C – 30 s and 72 °C – 30 s; finally 72 °C – 5 min. PCR products (expected size of 600 bp) were checked on 1.2% agarose gels and without purification used for third-stage PCR with primers MIDh-454-F and MIDh-454-R, containing barcodes (MIDx) and adapter sequences for sequencing on the 454 platform . Adapter sequences are described in matK marker was amplified using the primers matK_f (KIM3F) and matK_r (KIM1R) (https://www.qiagenbioinformatics.com).The  (KIM1R) . A totalmatK and 65 rbcL sequences generated in the present study, some published sequences , Bayesian inference (BI) and maximum likelihood (ML). The MP analyses were performed applying the parsimony ratchet (JK) support values were calculated in PAUP with 10,000 jackknife replicates and the TBR branch swapping algorithm with 36.788% of characters deleted and one tree held during each replicate. The ML analyses were done with RaxML  + Γ model as the standard predefined substitution model in RAxML. The BI analyses were performed with the MPI version of MrBayes  model space in the Bayesian MCMC analysis was applied (ML) and two partitions in the plastid DNA dataset . Two simultaneous runs of four parallel chains each were performed for 3 × 107 generations with a sample frequency of 1 tree per 2000 generations. Convergence of the runs was checked by making sure that the average standard deviation of split frequencies of the post-burn-in runs was below 0.01 and the effective sampling size (ESS) well above 200 in either run for all parameters. TreeGraph v.2 , Koelpinia (3 sp.), Pterachaenia (1 sp.), Epilasia (3 sp.) and Tourneuxia (1 sp.) were carpologically studied. Carpological data for the genera Tragopogon and Geropogon were taken from a previous study by In total, 149 species of oC. The sections were stained with 2% carbol fuchsin, then with 0.4% picroindigocarmine and 100% ethanol . The medium portion of the outer achenes is the most informative part, where all features are fully developed and was used for the carpological descriptions and phylogenetic reconstruction. Longitudinal sections were prepared in some cases to detect the peculiarities in the parenchymatous structures of the mesocarp. Prior to sectioning, the achenes were soaked in a mixture of ethyl alcohol, water and glycerol in equal proportions for a few days at 30–40The morphology of the achenes was documented using an Olympus SZ61 camera; the cross-sections were studied using Nikon Eclipse Ci microscopes and documented with a Nikon DS-Vi1 camera  at the Department of Higher Plants (Moscow State University). The ultrasculpture of the achene surface was investigated using an SEM (JSM-6380 LA) at the Laboratory of Electron Microscopy Center at Moscow M.V. Lomonosov State University.MP using the nrDNA tree as hypothesis of phylogenetic relationships and the ‘Trace character history’ and ‘Trace all characters’ tools as display methods. The character definitions and their state coding are provided under Results.Describing carpological features, we applied the tripartite descriptive model . The 50% majority consensus tree was largely congruent in topology with the 50% majority consensus trees of the BI and ML analyses. Fig. BI cladogram with the MP jackknife (JK) support values above and the BI posterior probabilities (PP) and ML bootstrap (BS) support values below the branches.The alignment of the nrITS region had a length of 707 characters; together with the coded indels, the matrix included a total of 832 characters, of which 422 were parsimony-informative. The Scorzonerinae are resolved as monophyletic with strong statistical support . The subtribe includes three major clades , which received strong  or full statistical support . Their relationships to each other are unresolved. Clade 1A solely comprises the genus Tourneuxia. Clade 1B includes one part of Scorzonera s.l. We designate it as Gelasia clade, because it includes Scorzoneravillosa, which provides the type of Gelasia Cass., being the oldest available generic name for members of this clade. Clade 1C includes the large remainder of the subtribe separated in two well-supported subclades. One  comprises the genera Tragopogon, Geropogon and Epilasia; the other  is a polytomy of four clades: (1) Scorzoneradivaricata forming a clade of its own; (2) Pterachaenia with S.codringtonii as sister; (3) Koelpinia and a S.polyclada clade; (4) a large clade including both Scorzonera in the sense of its type S.humilis and Podospermum in one well supported subclade  and in a second moderately supported subclade  Pseudopodospermum and an extended Takhtajaniantha clade. Further details will be treated in the Discussion below.The Phylogenetic reconstruction based on the concatenated plastid DNA markersMP analysis resulted in 103 most parsimonious trees . The 50% majority consensus tree was largely congruent in topology with the 50% majority consensus trees of the BI and ML analyses. Fig. BI cladogram with the MP jackknife (JK) support values above and the BI posterior probabilities (PP) and ML bootstrap (BS) support values below the branches.The alignment of the two concatenated plastid DNA markers had a length of 1387 characters, of which 112 were parsimony-informative. The Scorzonerinae are resolved as monophyletic and Tourneuxia as sister to the remainder of the subtribe with strong support . The latter clade 2 is a polytomy: clade 2A with almost full support includes the Gelasia clade; clade 2B with strong support only in MP includes Geropogon, Tragopogon and the Scorzoneradivaricata clade; clade 2C with low support  includes Pterachaenia with S.codringtonii, the S.polyclada clade and Koelpinia; clade 2D includes with moderate support  Pseudopodospermum with Epilasia as sister; clade 2E includes a moderately supported  Takhtajaniantha clade and the moderately strong supported clade 2F includes both Scorzonera in the sense of the type and Podospermum.The Epilasia appears as sister to a clade combining Geropogon and Tragopogon in the nrITS tree, whereas as sister to Pseudopodospermum in the plastid DNA tree; (2) Scorzoneradivaricata is sister to the Tragopogon-Geropogon clade in the plastid tree, but nested in a polytomy in the nrITS tree without the latter clade; (3) the Scorzonerapurpurea-S.renzii clades are nested in the nrITS tree in a polytomy with S.angustifolia and the S.albicaulis clade, but in the plastid DNA tree, these two species are found in a polytomy with the Podospermum clade.Fewer of the deep nodes are resolved in the plastid DNA tree compared to the nrITS tree, but both reconstructions revealed largely the same major terminal clades. Statistically significant topological differences are few. They mainly concern the relationships of three clades: (1) Scorzonerinae are, in principle, homocarpic. Minor differentiations corresponding to the centripetal development of the florets and thus also achenes in the Asteraceae capitula may, however, occur. The marginal achenes are considered as the most representative type. Achene wall anatomy and morphology are fully developed in the middle third of the achene as in all Asteraceae. Correspondingly, all features refer to the middle third of the marginal achenes.The capitula of the Cichorieae and all Compositae) that may be barely noticeable. The principal vascular bundles of the achene wall can be seen often well below the principal ribs even if the latter are not clearly expressed. In the Scorzonerinae, each segment forms one main rib and, usually, two secondary ribs, the latter shared with the contiguous segments, resulting in a pattern of 10 ribs, either differentiated or not into more prominent principal and less prominent secondary ribs. The achene epidermis is the outer cell layer of the achene wall and can be glabrous, hairy or papillose. The achene surface can also exhibit emergences: stout cell conglomerations, hooked spines or tubercles with verrucose ribs consisting of pore parenchyma. Below the epidermis, the achene wall is composed of sclerenchymatous and parenchymatous layers of various arrangements and thickness. Sometimes, air cavities are present, resulting from rupture of thin-walled parenchyma. The endocarp is often obliterated. The seed coat is ± adjoining to the endocarp and consists of two or several layers, sometimes with the vestiges of vascular bundles. The endosperm is two-layered and the vertical embryo occupies almost all volume in the achene corpus.The achene wall Fig.  is compoThe morphological characters of the pappus, as part of the diaspore, are also included in the carpological analysis.The carpologial analysis across the subtribe revealed a variety of features, which were found to be diagnostic for species or groups of them. In the following, these carpological features are coded in characters (composed of a structure and a property) and states. In the character designation, structures and substructures are separated by colons and these are separated from the respective property by a semicolon.1. Achene: carpopodium ; presence0: absent1: presentScorzonera s.l. .Carpopodia as a protrusion of the basal achene wall surrounding the stipe-like structure in which the vascular bundles of the achene enter the receptacle, are known throughout the family and the tribe . A parti2. Achene: carpopodium: border (by shape and surface texture) between achene corpus and carpopodium; presence0: absent Fig. 1: present Fig. Scorzonerasubg.Podospermum and some members of S.subg.PseudopodospermumWe note here for the first time that the carpopodium is morphologically separated from the achene corpus by a border in 3. Achene epidermis: hairs; presence0: none, epidermis glabrous1: with soft multicellular eglandular hairs Fig. 2: with drastically elongated, often slightly curved papillae looking like glandular hairs Fig. 3: with cylindrical, not drastically elongated papillae or mamillae Fig. 4: with retrorse acute papillae Fig. Koelpinia and Epilasia, respectively.The outgrowths , if present, originate from the achene epidermis. The hairs are multicellular in contrast to papillae  and mamillae  that are always unicellular. The character states 2 and 4 are peculiar for 4. Achene surface: emergences; presence0: absent1: present5. Achene surface: emergences; shape  equal in thickness present1: Discontinuous sclerenchymatous layers with a gap in the principal ribs2: Continuous sclerenchymatous layers with a narrow or wider hunch on either side of the principal rib3: Continuous or slightly discontinuous sclerenchyma with well-expressed invaginations (depressions).Sclerenchyma as stabilising element develops in the achene wall above the vascular bundles into different patterns and formations as defined in the states coding.7. Achene wall: sclerenchyma; orientation0: Sclerenchyma entirely with parallel orientation to achene axis1: Sclerenchyma differentiated: outer layer with parallel, inner with oblique to perpendicular orientation to achene axis.2: Sclerenchyma only in emergences or rib areas perpendicular (or oblique) to achene axis.8. Achene wall: parenchyma; arrangement Fig. 0: Present as subepidermal continuous layer(s)1: Continuous parenchyma layers above and below the sclerenchyma2: Insular in principal ribs below sclerenchyma and in secondary ribs above sclerenchyma3: Only insular in principal ribs below sclerenchyma4: Insular above sclerenchymatous invaginations and below sclerenchyma in the principal ribs5: Only insular above sclerenchymatous invaginations6: Absent7: Parenchyma continuous above sclerenchyma and below sclerenchyma insular in principal ribs8: Only insular in second ribs above sclerenchymaBesides sclerenchyma, the main element of the achene wall is parenchyma, but often the parenchyma elements are left behind in quantity by the sclerenchyma. The distribution of the parenchyma elements in the achene wall is taxon specific and several different arrangements have been found as outlined in the character state coding. The various ways the parenchyma can be arranged in the achene wall is schematically shown in Fig. 9. Achene wall: parenchyma; differentiation0: Parenchyma only as subepidermal mechanical parenchyma1: Subepidermal collenchyma-like layers differentiated besides parenchyma2: Parenchyma of thin-walled cells only3: Parenchyma of two types present, mechanical parenchyma and such with thin-walled cellsThe unusual collenchyma-like layers are built of thin-walled cells with prominent intercellular spaces of triangular shape. The parenchyma in character states 0 and 2–3 lack prominent intercellular spaces.10. Achene wall: air cavities; occurrence0: Absent1: Present11. Achene wall; ribbing pattern0: Each segment with a principal rib and 2 secondary ribs, the latter shared with the contiguous segments, achene middle third, thus with 5 principal and 5 secondary ribs1: Each segment with a principal rib only, achene middle third, thus with 5 ribs2: No distinct ribs developed, middle third ± terete (roundish)3: Two or more ribs enlarged to wings12. Achene wall: tannins; presence0: Absent1: Present in cell protoplast2: Present in cell walls only13. Achene beak; presence0: Absent1: PresentThe beak is defined as the more or less abrupt attenuation of the achene apex into a stipe-like structure carrying the pappus disc.14. Pappus; presence0: Absent1: Present15. Pappus; structure0: Of entirely softly plumose bristles 1: Of entirely softly plumose bristles and scabrid awns2: Of bristles softly plumose in lower and scabrid in upper half or third 4: Bristles scabrid completely or for the most partCichorieae consists of bristles with lateral projection (= fimbriae) not or little exceeding the diameter of the bristles, which make the bristles rough, thus “scabrid”. Alternatively, the lateral projection may be many times longer than the bristle diameter and the bristles thus featherlike or “plumose”. Plumose pappi in the Cichorieae have evolved as three different types, each specific for one subtribe 3: yellowish4: grey or blackishThe matrix of the 16 carpological characters is given in Table Tourneuxia cladeFig. 7A, BAchenes without carpopodium, with two ribs elongated into small wings; achene epidermis glabrous; emergences absent; subepidermal parenchyma almost continuous, more prominent in the winged areas and much thinner (1–3 layers) between them, of thin-walled cells only; sclerenchyma continuous, its fibres of parallel orientation, sometimes slightly obliquely orientated in the region of the wings; no air cavities; tannins absent.Gelasia cladeFig. 7C, DAchenes without carpopodium; achene epidermis often densely covered with soft multicellular hairs making a long woolly indumentum; emergences mostly absent; subepidermal parenchyma usually insular, of thin-walled cells only; sclerenchyma continuous with an invagination on either side of the principal rib, parallel; no air cavities; tannins absent.Geropogon cladeFig. 8A, BAchenes without carpopodium; achene epidermis with papillae, emergences present; subepidermal parenchyma continuous of thin-walled cells only; sclerenchyma continuous, parallel; air cavities present; tannins present.Tragopogon cladeAchenes without capopodium; achene epidermis usually with papillae; emergences absent or present; subepidermal parenchyma continuous, of thick-walled or thick- and thin-walled cells; sclerenchyma continuous, parallel; air cavities usually present; tannins absent.Epilasia cladeFig. 8C, DAchenes with carpopodium; achene wall with hair-like papillae; emergences absent; subepidermal parenchyma continuous, of thick-walled cells only; sclerenchyma continuous, parallel; air cavities absent; tannins present.Scorzoneradivaricata cladeFig. 9A, BAchenes without carpopodium; achene epidermis with papillae; emergences absent; subepidermal parenchyma of both thin- and thick-walled cells; sclerenchyma continuous, parallel; air cavities absent; tannins present.Pterachaenia cladeFig. 9C, DPterachaeniastewartii) or without (Pterachaeniacodringtonii); achene epidermis with papillae or glabrous; emergences absent; subepidermal parenchyma discontinuous, located above sclerenchyma between the principal ribs and below sclerenchyma in the rib areas or only insular above sclerenchyma, of both thin- and thick-walled cells; sclerenchyma continuous, parallel; air cavities absent; tannins absent.Achenes without carpopodium; with 2–3 elongated ribs forming wings , with an invagination on either side of the principal rib, parallel; air cavities absent; tannins usually absent .Koelpinia cladeFig. 10C, DK.tenuissima); emergences present; subepidermal parenchyma continuous, of thick-walled cells only; sclerenchyma continuous, parallel (but perpendicular in emergences); air cavities absent; tannins absent.Achenes without carpopodium; achene epidermis glandular-like (elongated) papillae  with parenchyma of both thin- and thick-walled cells and sclerenchyma continuous with a narrow or wider hunch on either side of the principal ribs  with a narrow or wider hunch on either side of the principal ribs  or absent; parenchyma insular in principal ribs below sclerenchyma and in secondary ribs above sclerenchyma, of thin-wall cells only; sclerenchyma continuous, parallel; air cavities absent; tannins absent.Achenes without carpopodium; achene epidermis with papillae; emergences present  subepidermal parenchyma represented by collenchymatous cells and sclerenchyma with a gap in the principal ribs ; air cavities absent; tannins absent.The results of the ancestral character reconstruction, based on the nrITS tree, are presented in Suppl. material Scorzonera is polyphyletic in all traditional circumscriptions, based on morphological data. Our broader sampling, however, elucidates an even higher extent of polyphyly than assumed so far. Actually, Scorzonera s.l. comprises at least six independent lineages.The molecular phylogenetic study, presented here, is the one with the broadest sampling across the subtribe to date and the first comparing reconstructions, based on nuclear ribosomal and plastid DNA markers. It fully confirms previous studies  in that Scorzonerinae are not resolved in either reconstruction and in previous analyses. However, our nuclear and plastid DNA trees are topologically largely congruent and supplement each other to some extent, allowing a first hypothesis on the major lineages of the Scorzonerinae and their relationships, which are discussed following the structure of the ITS tree  the Gelasia clade and (3) the remainder. In our plastid DNA tree  clade or the Gelasia and Tourneuxia clades as successive sisters to the remainder of the subtribe, but without statistical support.The nrITS tree Fig.  separatea clade,  the GelaTourneuxia lineage corroborates this region being part of the ancestral area of the tribe Cichorieae  leaves. The pollen structure of T.variifolia, consisting of 9  lacunae, is not unique amongst Scorzonerinae and found also in Pterachaenia and different groups of Scorzonera s.l. , it is resolved far remote from the core of Scorzonera in both our trees. This illustrates a surprisingly strong discrepancy between the traditional classification of the genus and molecular phylogenetic results. The lineage has already been resolved by Lasiospora clade”, after Scorzonerasubg.Lasiospora (Cass.) Tzvelev and the genus Lasiospora Cass.  shifts to glabrous achenes occur and (b) variously hairy achenes have been developed in three other clades of the subtribe: in the Podospermum clade, Pseudopodospermum  and the Takhtajaniantha clade .The olyclada , 1964, wis x = 7 . CarpoloPterachaenia lineage was first established as Scorzonerasect.Pterachaenia by Bentham  and habit, a similar achene anatomy apart from the wings in P.stewartii and a similar distribution area. However, following the literature, they differ in pollen structure .The ved Fig. . In the nia Fig. . In bothKoelpinia lineage includes only the small genus Koelpinia  with an Irano-Turanian distribution, extending into the S Mediterranean area. The plastid DNA tree places Koelpinia in closer relationship to both the S.polyclada and Pterachaenia lineages (with only moderate support) and the nrITS tree only to the former (but with strong support). A closer relationship between Koelpinia and Pterachaenia has already been revealed by Tragopogon  and both in turn are sister to the Scorzonera clade with fairly strong support .Its three major clades, which include the vast majority of Takhtajaniantha lineage encompasses the species of some sections of the typical subgenus of Scorzonera: S.sect.Fibrillosae , S.sect.Egregiae (S.tau-saghyz), S.sect.Parviflorae p.p. (S.mongolica), S.sect.Papposae p.min.p. (S.capito), S.sect.Polyclada p.min.p. (S.pseudodivaricata); it also includes S.pusilla, the only member of S.sect.Pusillae, which was split from Scorzonera as the monotypic genus Takhtajaniantha by Scorzonera. The basic number of x = 7 is also shared by the other members of the lineage so far known. Later, T.pusilla, having only six abporal lacunae, to S.austriaca, S.ikonnikovii and S.tau-saghyz, which is a pollen type, however, also present in the Scorzonera s.str. clade   and into Central Europe and South Siberia (S.austriaca).The r. clade . TakhtajPseudopodospermum lineage has full support in the nrITS tree and moderately strong support  in the plastid DNA tree. The generic rank proposed by S.subg.Pseudopodospermum, which is typified by Scorzoneramollis, is corroborated by the fact that this lineage is resolved in both our reconstructions, separate from the core of Scorzonera. The Pseudopodospermum lineage, however, also includes species previously placed in sections Incisae, Foliosae, Papposae and Hissaricaeof the formersubgenusScorzonera . All members share a basic chromosome number of x = 7. It includes four major terminal clades: the Scorzonera s.str. clade, the S.purpurea clade, the S.albicaulis clade and the Podospermum clade. Besides, S.rupicola, an intricately branched shrublet and S.renzii, a linear-leafy perennial herb with peculiar involucre, both from Iran and Turkey, respectively, form separate clades. The latter, little known species was previously placed in S.sect.Turkestanicae , S.aristata and S.parviflora. The close relationship of S.humilis with S.aristata and S.parviflora was assumed already by S.sect.Parviflorae by S.mongolica by S.aristata; Scorzoneraparviflora is distributed from Central Europe to Central Asia, whereas S.humilis is widespread in Europe and S.aristata restricted to SW Europe : Scorzonerapurpurea and its close relative S.rosea were often considered to belong to S.subg.Podospermum due to the presence of a carpopodium (Takhtajaniantha), combined with often graminoid leaves and purple or lilac florets, pollen with 18  lacunae, narrow achenes with the same carpological type (present investigation). Carpology corroborates the close affinity of S.rhodantha (SE Europe) to this clade and it has sometimes been considered as subspecies of S.purpurea .Scorzoneraalbicaulis clade: This clade includes species of Scorzonerasect.Piptopogon, S.sect.Turkestanicae and S.sect.Polycladae. Morphologically, all species share the graminoid leaves, achenes attenuated into a more or less prominent beak, an easily caducous pappus and an achene surface slightly scabrid due to attenuate elongations of the epidermis cells. The presence of beaked achenes was the reason for Achyroseris Sch.Bip., based on A.macrospermum Sch.Bip.  and for the transfer of further species with beaked achenes from Scorzonera to Achyroseris by S.albicaulis clade are predominantly distributed in Central Asia, extending to W Asia and the N Himalaya (S.virgata); one species (S.angustifolia) is confined to the Iberian Peninsula and Morocco. They share a pollen with 24  lacunae  have the same achene anatomy as the Scorzonera s.str. clade . Carpological and morphological characters suggest the close affinity of S.crassicaulis, S.franchetii, S.petrovii, S.rupicola (not studied by molecular phylogeny) to this clade. lacunae . Carpolo lacunae  in S.anade Fig. , but oftPodospermum clade: This clade unites the majority of the members of Scorzonerasubg.Podospermum sensu Podospermum sensu S.hieraciifolia, S.songorica). Moreover, the pinnately lobed leaves are shown in our analysis to be a homoplastic state, because the pinnately leafy species of the former Podospermum sections Incisae  and Brevicaulis (S.brevicaulis) are resolved as members of the Pseudopodospermum clade. All members of the Podospermum clade have a well-developed carpopodium, which is another homoplasy shared with the Pseudopodospermum clade and also with the S.purpurea clade. Peculiar for the members of the Podospermum clade are: (1) horn-like appendages on the outer phyllaries of the capitula and (2) the diversely orientated sclerenchymatous fibres in the mesocarp of the achene wall. A limited number of species, investigated palynologically, share pollen with 24  lacunae (S.songorica), N Africa and S and Central Europe (S.laciniata). lacunae . Most spScorzoneralaciniata and S.cana occupy different positions in both the nrITS and plastid DNA trees  and the length of the ligules in relation to the involucre.Resolution within the clade is very poor and also, morphologically, its members are fairly uniform, indicating a young diversification age. Some samples of the widely distributed, variable species ees Figs , 2. ThisAccording to the ancestral character reconstruction based on the nrITS tree Fig. , achenesTourneuxia matches this ancestral type most closely, except for the presence of ribs enlarged to wings .Scorzonerinae are the rare exception  and stout hair-like papillae (Epilasia).The plesiomorphic state of an achene surface with unicellular papillae or mamillae . In other members of these two clades and in the Gelasia clade, the sclerenchyma shows well-expressed depressions. A differentiation of the sclerenchyma orientation with the outer layers parallel and the inner perpendicular to the achene axis seems a non-homoplastic synapomorphy of the Podospermum clade  are present in the Scorzonera clade; the distribution of this state in the Podospermum clade for which otherwise an arrangement continuous above and insular below the sclerenchyma (8/7) was revealed as synapomorphy, indicates a close relationship between these two states. Regarding its differentiation, the parenchyma has apparently been differentiated several times in the evolution of the subtribe to support stabilisation of the achene wall, either as mechanical or collenchyma-like tissue or both (9/1–3).The parenchyma of the achene wall shows considerable diversity regarding arrangement  as an exclusive synapomorphy of in the Pseudopodospermum clade .Occurrence of tannins  in Geropogon, the opposite shift to purely plumose bristles (15/0) in species of Tragopogon. With respect to pappus colour . Shifts are indicated to have occurred to a dirty white colour at some state of uncertain position in the evolution of the subtribe, with further shifts to yellowish, pure white or grey and reversals to fulvous.The pappus has been lost once in the Scorzonerinae are often not well distinguished morphologically. Neither gross morphology nor fruit anatomy provides non-homoplastic synapomorphies for most of the major lineages. A prominent example is Gelasia, of which many species are nicely recognisable by the very conspicuous long-lanate achene indumentum; in a number of species, however, a reversal to glabrous achenes has occurred and what remains is some overall similarity of the Gelasia species which cannot be appropriately expressed in a character-state matrix or an identification key. This situation is apparently responsible for the reluctant reception of any classification of Scorzonerinae with more than the few morphologically conspicuous elements and the perseverance of a polyphyletic taxonomic concept of Scorzonera in spite of contrary evidence. We assume, however, that practical taxonomic experience in the application of a phylogenetic classification will bring to light new means to distinguish the various entities.We have shown above that morphology, even extended to include fruit anatomy, does not very well reflect the structure of the subtribe as revealed through molecular phylogenetics. Actually, most of the lineages resolved are difficult to characterise by morphology. Even more distant clades of the Gelasia clade and retain the remainder of the current Scorzonera, but also include in it the genera Pterachaenia and Koelpinia. In this option Scorzonera would encompass clade 1C2  which is definitely undesirable – and consequently shared by all three available options. The second option is to recognise Scorzonera in the circumscription of clade 2  recognised, with nomenclature (typifications and synonymies), diagnostic features, brief descriptions, species lists  and the distribution area  lacunae  between them, of thin-walled cells only, air cavities absent, tannins absent.Pappus: 4–7 mm, obliquely inserted in the marginal achenes, bristles plumose almost entirely, but scabrid in upper portion, the fimbriae of which are soft and tangled with each other, proximally brownish, distally dirty white.x = 7, diploid.T.variifolia Coss. – Fig. 15A, B(1) N Africa .Taxon classificationPlantaeAsteralesAsteraceaeCass., Bull. Sci. Soc. Philom. Paris 1818: 33. 1818.B646E85A-3172-530D-A1B8-E8D6DDED42F8Scorzonerasect.Gelasia (Cass.) DC., Prodr. 7(1): 123. 1838. Type: Scorzoneravillosa Scop. ≡ Gelasiavillosa (Scop.) Cass.  ≡ Lasiospora Cass. in Cuvier, Dict. Sci. Nat. 25: 306. 1822.  = Scorzonerasect.Lasiospora (Cass.) Less., Syn. Gen. Compos.: 134. 1832.  ≡ LasiosporaScorzonerasubg.Lasiospora (Cass.) Peterm., Deutschl. Fl.: 334. 1846–1849. Lectotype (Lasiosporahirsuta (Gouan) Cass.  ≡ ectotype : 45: LasScorzonerasubsect.Pulvinares Boiss., Fl. Orient. 3: 756. 1875.  = Scorzonerasect.Pulvinares (Boiss.) Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 24. 1935. Lectotype (designated here): Scorzoneraseidlitzii Boiss.  ≡ Scorzonerasubsect.Infrarosulares Hand.-Mazz., Ann. K. K. Naturhist. Hofmus. 27: 455. 1913.  = Scorzonerasect.Infrarosulares (Hand.-Mazz.) Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 10. 1935. Lectotype (designated here): Scorzoneraacantholimon Hand.-Mazz.  ≡ Scorzonerasect.Nervosae Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 126. 1935. Type: Scorzoneralatifolia (Fisch. & C.A.Mey.) DC.  = Scorzonerasect.Trachyactis Rech.f., Oesterr. Bot. Z. 84: 169. 1935. Lectotype (designated here): Scorzoneracretica Willd. Note: This section was described, based on the entirely scabrid pappus, a remarkable feature in Gelasia and the subtribe. Three species were reported to form this section: S.dependens Rech.f. (= S.cretica Willd.), S.araneosa Sm. and S.eximia Rech.f. The first species (as S.cretica) was nested in our nrITS phylogeny in the Gelasiaclade. Two species, S.cretica and S.araneosa, were included in our carpological analysis and match the carpology of Gelasia.  = Scorzonerasect.Tuberosae Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 39. 1939. Type : Scorzoneraensifolia M.Bieb.  = Scorzonerasect.Anatolia Makbul & Coşkunç., Turk. J. Bot. 39: 77. 2015. Type: S.zorkunensis Coskunç. & Makbul  = Perennials; leaves entire; involucre often pubescent; achenes often densely lanate, rarely glabrous, usually smooth rarely with emergence; pappus often rigid, plumose in lower portion and scabrid in upper portion or rarely almost completely scabrid; sclerenchyma forming a sheath, with an invagination on either side of the principal rib.Habit, life form, subterranean parts: perennial herbs of various habits, rarely subshrubs (only G.acantholimon), with a taproot, more rarely with globose or cylindrical tuber.Leaves: rosulate or cauline, sessile or petiolate, linear to ovate, often hairy  or glabrous, margin entire and flat or undulate.Stem, synflorescence: scape-like, small or leafy, tall and branched, often pubescent, more rarely glabrous , synflorescence racemiform, spiciform or corymbiform or capitula single.Capitula: involucre usually pubescent, sometimes glabrescent at fruiting or rarely glabrous, phyllaries in several series, all lanceolate or triangular, outer phyllaries < 3–4 times smaller than the inner ones, receptacle glabrous or hairy, capitula of many yellow florets , well exceeding the involucre.ets Fig.  or, rarePollen: echinolophate, tricolporate and each colpus divided into 2 lacunae; with 18  or, more rarely, 24  lacunae , parenchyma usually insular above invaginations of the sclerenchyma and below the principal ribs or only insular above invaginations of sclerenchyma, air cavities absent, sclerenchymatous layers continuous or discontinuous in the main ribs, forming a sheath, with an invagination on either side of the principal rib, fibres orientated parallel to the fruit axis, tannins absent.Pappus: 5–24 mm; bristles plumose in lower portion and scabrid in upper portion or more rarely, almost completely scabrid; dirty white, yellow or fulvous.x = 6 (rarely 7), diploids or tetraploid.Gelasiaacantholimon (Hand.-Mazz.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraacantholimon Hand.-Mazz. in Ann. K. K. Naturhist. Hofmus. 27: 455. 1913. urn:lsid:ipni.org:names:77204005-1(1) Gelasiaalbicans (Coss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraalbicans Coss., Notes Crit.: 11. 1851. urn:lsid:ipni.org:names:77204006-1(2) Gelasiaaraneosa (Sm.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraaraneosa Sm. in Sibthorp, Fl. Graec. Prodr. 2(1): 123. 1813. urn:lsid:ipni.org:names:77204007-1(3) Gelasiaaucheriana (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraaucheriana DC., Prodr. 7(1): 125. 1838. urn:lsid:ipni.org:names:77204009-1(4) Gelasiabiebersteinii (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerabiebersteinii Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 95. 1939. urn:lsid:ipni.org:names:77204011-1(5) Gelasiacaespitosa (Pomel) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracaespitosa Pomel, Nouv. Mat. Fl. Atl.: 266. 1875. urn:lsid:ipni.org:names:77204012-1(6) Gelasiacallosa (Moris) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracallosa Moris, Stirp. Sard. Elench. 1: 29. 1827. urn:lsid:ipni.org:names:77204014-1(7) Gelasiacinerea (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracinerea Boiss., Diagn. Pl. Orient., ser. 1, 11: 44. 1849. urn:lsid:ipni.org:names:77204016-1(8) Gelasiacircumflexa (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracircumflexa Krasch. & Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., ser. 2, 42. 148. 1934. urn:lsid:ipni.org:names:77204017-1(9) Gelasiacretica (Willd.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracretica Willd., Sp. Pl. 3: 1504. 1803. urn:lsid:ipni.org:names:77204018-1(10) Gelasiadoriae  Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneradoriae Degen & Bald., Oesterr. Bot. Z. 46: 417. 1896. urn:lsid:ipni.org:names:77204021-1(11) Gelasiadzhawakhetica (Sosn. ex Grossh.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneradzhawakhetica Sosn. ex Grossh., Fl. Kavk. 4: 236. 1934. urn:lsid:ipni.org:names:77204022-1(12) Gelasiaensifolia (M.Bieb.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraensifolia M.Bieb., Fl. Taur.-Caucas. 2: 235. 1808. urn:lsid:ipni.org:names:77204023-1(13) Gelasiaeriophora (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraeriophora DC., Prodr. 7(1): 125. 1838. urn:lsid:ipni.org:names:77204025-1(14) Gelasiafilifolia (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerafilifolia Boiss., Fl. Orient. 3: 774. 1875. urn:lsid:ipni.org:names:77204026-1(15) Gelasiahirsuta (Gouan) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Tragopogonhirsutus Gouan, Fl. Monsp.: 342. 1764. urn:lsid:ipni.org:names:77204027-1(16) Gelasiaketzkhovelii (Grossh.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraketzkhovelii Sosn., Trudy Tbilissk. Bot. Inst. 2: 219. 1938. urn:lsid:ipni.org:names:77204029-1(17) Gelasiakotschyi (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerakotschyi Boiss., Fl. Orient. 3: 780. 1875. urn:lsid:ipni.org:names:77204031-1(18) Gelasialanata (L.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Leontodonlanatus L., Cent. Pl. 1: 26. 1755. urn:lsid:ipni.org:names:77204032-1(19) Gelasialasiocarpa (Chamberlain) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneralasiocarpa Chamberlain, Notes Roy. Bot. Gard. Edinburgh 33: 255. 1974. urn:lsid:ipni.org:names:77204034-1(20) Gelasialatifolia (Fisch. & C.A.Mey.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Lasiosporalatifolia Fisch. & C.A.Mey., Index Sem. Hort. Petrop. 1: 30. 1835. urn:lsid:ipni.org:names:77204035-1(21) Gelasialitwinowii (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneralitwinowii Krasch. & Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., ser. 2, 43: 153. 1934. urn:lsid:ipni.org:names:77204036-1(22) Gelasialongiana (Sümbül) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneralongiana Sümbül, Edinburgh J. Bot. 48(1): 35. 1991. urn:lsid:ipni.org:names:77204039-1(23) Gelasiamackmeliana (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneramackmeliana Boiss., Diagn. Pl. Orient., ser. 1, 11: 44. 1849. urn:lsid:ipni.org:names:77204041-1(24) Gelasiamirabilis (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneramirabilis Lipsch., Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 22: 288. 1963. urn:lsid:ipni.org:names:77204043-1(25) Gelasiapisidica (Hub.-Mor.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapisidica Hub.-Mor., Bauhinia 7(3): 179. 1982. urn:lsid:ipni.org:names:77204044-1(26) Gelasiapsychrophila (Boiss. & Hausskn.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapsychrophila Boiss. & Hausskn. in Boissier, Fl. Orient. 3: 777. 1875. urn:lsid:ipni.org:names:77204045-1(27) Gelasiapygmaea (Sm.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapygmaea Sm., Fl. Graec. Prodr. 2(1): 123. 1813. urn:lsid:ipni.org:names:77204046-1(28) Gelasiaramosissima (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraramosissima DC., Prodr. 7(1): 125. 1838. urn:lsid:ipni.org:names:77204047-1(29) Gelasiarigida (Aucher ex DC.) Zaika, Sukhor. & N.Kilian, comb. nov. : 123. 1838. urn:lsid:ipni.org:names:77204048-1(30) ov. Fig.  ≡ ScorzoGelasiasandrasica (Hartvig & Strid) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerasandrasica Hartvig & Strid, Bot. Jahrb. Syst. 108(2–3): 311. 1987. urn:lsid:ipni.org:names:77204050-1(31) Gelasiaseidlitzii (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraseidlitzii Boiss., Fl. Orient. 3: 775. 1875. urn:lsid:ipni.org:names:77204051-1(32) Gelasiasericea (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerasericea DC., Prodr. 7(1): 123. 1838. urn:lsid:ipni.org:names:77204053-1(33) Gelasiatomentosa (L.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneratomentosa L., Sp. Pl., ed. 2, 2: 1112. 1763. urn:lsid:ipni.org:names:77204054-1(34) Gelasiatuberosa  Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneratuberosa Pall., Reise Russ. Reich. 3: 757. 1776. urn:lsid:ipni.org:names:77204057-1(35) Gelasiaulrichii (Parolly & N.Kilian) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraulrichii Parolly & N.Kilian, Willdenowia 32: 198. 2002. urn:lsid:ipni.org:names:77204058-1(36) Gelasiavillosa (Scop.) Cass.(37) N Africa: DZ; EG; LY; MA; TN. Europe: AL; BA; CY; ES; FR; GR; HR; IT; MD; ME; RS; RU; SI; UA. Asia-Temperate: AF; AM; AZ; CH; GE; IL; IQ; IR; JO; KG; KZ; LB; RU (Asiatic part); SY; TJ; TM; UZ. Asia-Tropical: PK.Gelasia, based on their morphological characters : Scorzoneraamasiana Hausskn. & Bornm., S.boissieri Lipsch., S.bungei Krasch. & Lipsch., S.charadzae Papava, S.gageoides Boiss., S.ispahanica Boiss. ≡ Lasiosporaispahanica (Boiss.) Walp., S.joharchii S.R.Safavi, S.karabelensis Parolly & N.Kilian, S.kozlowskyi Sosn. ex Grossh., S.lipskyi Lipsch., S.persica Boiss. & Buhse, S.pulchra Lomak., S.safievii Grossh., S.sahnea Parsa, S.scopariiformis Lipsch., S.sublanata Lipsch., S.sericeolanata (Bunge) Krasch. & Lipsch., S.veratrifolia Fenzl ≡ Lasiosporaveratrifolia (Fenzl) Walp., S.wendelboi Rech.f., S.woronowii Krasch., S.xylobasis Rech.f., S.yildirimlii A.Duran & Hamzaoğlu, S.zorkunensis Coşkunç. & MakbulThe following species may also belong to Taxon classificationPlantaeAsteralesAsteraceae(Bunge) Benth. in Bentham & Hooker, Gen. Pl. 2: 532. 1873.0D02DDA2-9E5C-5225-ACDD-B2F928FF5430Scorzonerasect.Epilasia Bunge, Beitr. Fl. Russl.: 200. 1852. Lectotype (designated by Epilasiahemilasia (Bunge) Kuntze  ≡ Annual herbs; phyllaries in two rows, outer herbaceous, leaflike, about equalling in length the inner phyllaries; pollen with 18 lacunae; achenes with carpopodium; pappus arising from flat or caplike pappus disk, grey to blackish; bristles plumose, 5–10 of them longer, scabrid distally.Habit, life form, subterranean parts: annual herbs with taproot.Leaves: rosulate and lower stem leaves petiolate, upper stem leaves sessile, lanceolate to ovate, margin flat or somewhat undulate and serrulate.Stem, synflorescence: stems solitary or more frequently several (three to five), glabrous to arachnoid hairy; capitula terminal, solitary or up to eight on the stem.Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae, with 18  lacunae  or with conic caplike pappus disk and pappus covering the upper half of the achene (E.hemilasia); pericarp with thin-walled subepidermal parenchymatous sheath, the cell walls of which can be filled with the tannins and continuous sclerenchymatous layers orientated parallel to the fruit axis (but sometimes perpendicular in the ribs of E.mirabilis), air cavities absent.Pappus: 5–8 mm, dense, ash-grey or rusty, five of its bristles (rarely more) fragile, barbellate at tip, other bristles long-plumose or rarely bristles plumose in lower portion and scabrid in upper portion.x = 12, amphiploids.E.acrolasia (Bunge) C.B.Clarke ex Lipsch.(1) E.hemilasia (Bunge) Kuntze(2) E.mirabilis Lipsch.(3) Asia-Temperate: AF; AM; AZ; CN; GE; IQ; IR; KG; KZ; LB; SY; TJ; TM; UZ. Asia-Tropical: PK.Taxon classificationPlantaeAsteralesAsteraceaeL., Sp. Pl. 2: 789. 1753.B5EC8B81-0647-56F5-B1FE-ADE1B093FF74Tragopogonporrifolius L., linear to oblong, margin entire or undulate.Stem, synflorescence: stem solitary or branched, always leafy, glabrous to lanate; peduncle sometimes inflated.Capitula: involucre glabrous to lanate, cylindrical, phyllaries in one series, triangular or oblong, equal in size, acute, receptacle flat, naked; with > 20 florets, of various length ratios compared to involucre, yellow, orange, violet or purple.Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae, with 15  lacunae  Sch.Bip.)Involucre with phyllaries in a single series; pappus of marginal achenes of five scabrid rays, pappus of inner achenes with plumose bristles.Habit, life form, subterranean parts: annual, ephemere.Leaves: rosulate and cauline, graminoid , linear or lanceolate, margin entire.Stem, synflorescence: stem solitary or branched, always leafy, glabrous; peduncle slightly inflated.Capitula: involucre glabrous, cylindrical, phyllaries 7–8, linear-lanceolate, in a single series, equal in size, to ~30 mm; receptacle flat, marginally hairy; florets 10–20, much exceeded by the involucre, violet or purple  lacunae  Sch.Bip.(1) N Africa: EG; DZ; LY; MA; TN. Europe: AL; BG; CY; ES; FR; GR; HR; IT; MD; MT; PT; RU; UA. Asia-Temperate: AM; AZ; IQ; IL; IR; JO; LB; SY; TR.Rare alien in DE.Taxon classificationPlantaeAsteralesAsteraceaeZaika, Sukhor. & N.Kiliangen. nov.9DC8F22F-99A0-5BB9-82EE-E42785969877urn:lsid:ipni.org:names:77204059-1Scorzonerasect.Polyclada DC., Prodr. 7(1): 125. 1838. Lectotype (designated here): Scorzoneradivaricata Turcz.  = Lipschitziadivaricata (Turcz.) Zaika, Sukhor. & N.Kilian.Scorzoneradivaricata clade in our phylogenetic analysis. We treat the species in its narrow sense, excluding, for the time being, the varieties with more numerous florets (7–12) per capitulum, because they are likely misplaced in this species .This new genus corresponds to the monotypic Subshrubs or perennial herbs; caudex with smooth scarious leaf sheath residues; stem leaves linear to filiform, only to 1 cm long, apically usually hooked; capitula numerous, with 4–5 florets.Habit, life form, subterranean parts: subshrubs or perennial herbs with branched caudex covered with smooth scarious leaf sheath residues.Leaves: sessile, linear to filiform, up to 1 cm, apically often hooked.Stem, synflorescence: stem divericately branched, glabrous; capitula many, terminal.Capitula: involucre puberulent, phyllaries in two series, outer phyllaries tiny, triangular or ovate, inner phyllaries oblong; receptacle naked, capitula with 4 or 5 florets; florets yellow, almost equal in length to the involucre.Pollen: data n/a.Achenes: 6–10 mm, cylindrical, with 10 ribs, smooth only apically with cylindrical papillae, without emergences and carpopodium; achene wall with both thin- and thick-walled cells  and continuous sclerenchymatous layers which cells orientated parallel to the achene axis, air cavities absent.Pappus: 5–8 mm, bristles plumose in lower part and scabrid in upper portion, dirty-white.Scorzoneradivaricata).x = 7, diploid (Scorzonera.The new genus is named after Sergey Yu. Lipschitz [Lipshits] (1905–1983), a Russian botanist and monographer of L.divaricata (Turcz.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneradivaricata Turcz., Bull. Soc. Imp. Naturalistes Moscou 5: 200. 1832. urn:lsid:ipni.org:names:77204060-1(1) Asia-Temperate: CN; MN.Taxon classificationPlantaeAsteralesAsteraceae(Benth.) Stewart, Punjab Forest Rec. 2, 1 [Cat. Pl. Rawalpindi Distr.]: 50. 1952.8BFED08C-C176-5B1A-8C15-61AA92B573DFScorzonerasect.Pterachaenia Benth. in Bentham & Hooker, Gen. Pl. 2: 532. 1873.Type: Pterachaeniastewartii (Hook.f.) R.R.Stewart  ≡ Pterachaenia. His statement “I think this section will also turn out to be a separate genus (gen. proprium Pterachaenia)” [translated from Russian] qualifies it, however, as a provisional genus name not formally accepted by its author.Usually Flowering stems several or many, unbranched, leafless (scapes); phyllaries lanceolate, acute; florets yellow with red veins; achenes with 2–3 wings or without.Habit, life form, subterranean parts: perennial with caudex or annual herbs, with taproot.Leaves: rosulate, graminoid, linear or lanceolate, glabrous or hairy, often broadened in upper half.Stem, synflorescence: stems leafless, unbranched (scapes) several to many, pubescent in lower part and glabrous in upper part, with terminal capitulum.Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae, with 9  or 18  lacunae  or without wings (P.codringtonii), papillate and with small emergences (spinulae) (P.stewartii) or with smooth surface (P.codringtonii), with five principal ribs, parenchyma of two types, with thick-walled cells and with thin-walled cells, discontinuous  or only insular above sclerenchyma, air cavities absent, sclerenchyma continuous, equal in thickness, fibres orientated parallel to the fruit axis, tannins absent.Pappus: 13–20 mm, fulvous, bristles plumose, apically scabrid.Pterachaeniastewartii) and x = 7 (P.codringtonii), diploids.x = 6  Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracodringtonii Rech.f., Oesterr. Bot. Z. 97: 263. 1950. urn:lsid:ipni.org:names:77204061-1(1) Pterachaeniastewartii (Hook.f.) R.R.Stewart in Punjab Forest Rec. 2, 1: 50. 1952.(2) Taxon classificationPlantaeAsteralesAsteraceaePall., Reise Russ. Reich. 3: 755. 1776.19EBB35B-CA75-5833-9C97-C8192AFB5695Koelpinialinearis Pall.K.tenuissima), without pappus, but apically with hooked, retrorse spines.Annual herbs; pollen with 15 lacunae; at least outer achenes columnar-scorpioid, surface with hooked emergences and elongated papillae looking like glandular hairs,  lacunae, often pollen tetracolporate in polyploids annular (K.macrantha) or almost straight (K.tenuissima and K.turanica), with 5 or 10 more or less prominent ribs, surface often covered with emergences resembling hooked spines and glandular-like (elongated) papillae or surface almost glabrous in K.tenuissima, without carpopodium; achene wall with or without air cavities, parenchyma continuous above sclerenchyma represented by thick-walled cells, sclerenchyma continuous, its fibres orientated parallel to the fruit axis (perpendicular in emergences), tannins absent.Pappus: bristles absent, achenes with several retrorse hooked spines  K.linearis Pall.(2) K.macrantha C.Winkl.(3) K.tenuissima Pavlov & Lipsch.(4) K.turanica Vassilcz.(5) N Africa: DZ; EG; LY; TN. Asia-Temperate: AF; AM; AZ; BH; CN; CY; GE; IL; IQ; IR; KG; KW; KZ; LB; QA; SA; SY; TJ; TM; TR; UZ. Asia-Tropical: IN; PK. Europe: ES; RU; UA.Taxon classificationPlantaeAsteralesAsteraceaeZaika, Sukhor. & N.Kiliangen. nov.008D20C2-FEB1-53F1-B00E-9969CEB94DD5urn:lsid:ipni.org:names:77204063-1Scorzonerasubsect.Intricatae Boiss., Fl. Orient. 3: 756. 1875.  = Scorzonerasect.Intricatae (Boiss.) Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 10. 1935. Type  Zaika, Sukhor. & N.KilianScorzonerapolyclada clade. Morphologically, Ramaliella resembles Lipschitziadivaricata, but differs by the caudex surface, size of the capitula and the achene anatomy of a type of its own.The new genus corresponds to the Subshrubs or perennial herbs; stems numerous, intricately and divaricately branched Fig. ; leaves Habit, life form, subterranean parts: subshrubs or perennial herbs with taproot, often with cushion-like habit.Leaves: few, the lower leaves filiform, the upper ones reduced, curved, entire or denticulate.Synflorescence: stems numerous, strongly divaricately and intricately branched, glabrous or puberulent, capitula terminal.Capitula: involucre glabrous or pubescent, phyllaries in several series, the outer phyllaries tiny, triangular, the inner phyllaries linear-lanceolate much longer than the outer ones, receptacle naked, flat, florets 3–12, yellow, slightly exceeding the involucre  lacunae  barely noticeable secondary ribs, surface smooth; achene wall with parenchyma of both thin- and thick-walled cells, sclerenchyma forming a sheath (sometimes irregularly discontinuous), with an invagination on either side of the principal rib, cells orientated parallel to the achene axis, air cavities absent, tannins absent or rarely present in cell wall only (R.koelpinioides).Pappus: 11–18 mm, fulvous or dirty white, bristles plumose, but in upper part scabrid.R.koelpinioides and R.tortuosissima: x = 7, diploid (data for The new genus is named after the specific “brushwood” habit (rāmālia: brushwood).Ramaliellaintricata (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraintricata Boiss., Diagn. Pl. Orient., ser. 1, 7: 9. 1846. urn:lsid:ipni.org:names:77204064-1(1) Ramaliellakoelpinioides (Rech.f.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerakoelpinioides Rech.f., Fl. Iranica 122: 53. 1977. urn:lsid:ipni.org:names:77204065-1(2) Ramaliellalongipapposa (Rech.f.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneralongipapposa Rech.f., Fl. Iranica 122: 53. 1977. urn:lsid:ipni.org:names:77204066-1(3) Ramaliellamusilii (Velen.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneramusilii Velen., Sitzungsber. Königl. Böhm. Ges. 9: 8. 1911. urn:lsid:ipni.org:names:77204068-1(4) Ramaliellapolyclada (Rech.f. & Köie) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapolyclada Rech.f. & Köie, Biol. Skr. 8(2): 195. 1955. urn:lsid:ipni.org:names:77204069-1(5) Ramaliellatortuosissima (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneratortuosissima Boiss., Fl. Orient. 3: 775. 1875. urn:lsid:ipni.org:names:77204071-1(6) N Africa: EG. Asia-Temperate: AE; AF; IQ; IR; JO; KU; SA; YE. Asia-Tropical: PK.Ramaliella, based on their morphological characters : Scorzonerahondae Kitam., S.microcalathia (Rech.f.) Rech.f., S.subaphylla Boiss., S.yemensis PodlechThe following species may belong to Taxon classificationPlantaeAsteralesAsteraceae(Lipsch. & Krasch.) Kuth., Kavkaz. Predstav. Scorzonerinae: 85. 1978.976E0259-90D6-5E0E-9120-F3BC0CD42C3FScorzonerasect.Pseudopodospermum Lipsch. & Krasch. in Lipschitz, Fragm. Monogr. Gen. Scorzonera 1: 70. 1935.  ≡ Scorzonerasubg.Pseudopodospermum (Lipsch. & Krasch.) Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 48. 1964. Lectotype  Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 721. 1964. Lectotype  Kuth., Kavkaz. Predstav. Scorzonerinae: 99. 1978. Type  amplexicaule, their shape diverse (linear-lanceolate to broadly ovate), margin entire and then often undulate, or dentate to pinnatisect.Stem, synflorescence: stem solitary or with a few branches, sometimes leafless and scapose, glabrous to lanate; capitula terminal.Capitula: involucre glabrous to lanate, phyllaries in several series, outer phyllaries triangular or ovate, obtuse, inner phyllaries oblong or lanceolate, much longer than outer phyllaries, receptacle naked, flat, florets more than 12, exceeding the involucre up to 1.5 times, yellow or purple  lacunae , often reduced to one- to several layers consisting of thin- and/or thick-walled cells, sometimes with subepidermal mechanical parenchyma, air cavities absent or present, sclerenchyma discontinuous with a gap in the principal ribs or continuous forming a sheath, equal in thickness or with a narrow or wider hunch on either side of the principal rib, sclerenchymatous fibres orientated parallel to the achene axis, tannins absent or present in the cell protoplasts.Pappus: 6–28 mm, bristles equal or unequal (5–10 bristles often longer than the rest), plumose in lower and scabrid in upper part, dirty white or white, yellowish, blackish or rarely fulvous.x = 7, diploids, tetraploids or hexaploids.Pseudopodospermumboeticum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneramontanavar.boetica Boiss. in Candolle, Prodr. 7(1): 121. 1838 ≡ Scorzoneraboetica (Boiss.) Boiss., Voy. Bot. Espagne 2: 382. 1841. urn:lsid:ipni.org:names:77204074-1(1) Pseudopodospermumbrevicaule (Vahl) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerabrevicaulis Vahl, Symb. Bot. 2: 88. 1791. urn:lsid:ipni.org:names:77204076-1(2) Pseudopodospermumcalyculatum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracalyculata Boiss., Diagn. Pl. Orient., ser. 1, 11: 42. 1849. urn:lsid:ipni.org:names:77204078-1(3) Pseudopodospermumchantavicum (Pavlov) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerachantavica Pavlov, Vestn. Akad. Nauk Kazakhsk. S.S.R. 8: 27. 1950. urn:lsid:ipni.org:names:77204079-1(4) Pseudopodospermumcrispatulum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerahispanicavar.crispatula DC., Prodr. 7(1): 121. 1838 ≡ Scorzoneracrispatula (DC.) Boiss., Voy. Bot. Espagne 2: 741. 1845. urn:lsid:ipni.org:names:77204082-1(5) Pseudopodospermumcrocifolium (Sm.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracrocifolia Sm. in Sibthorp, Fl. Graec. Prodr. 2(1): 123. 1813. urn:lsid:ipni.org:names:77204084-1(6) Pseudopodospermumdavisii (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneradavisii Lipsch., Bot. Mater. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 22: 291. 1963. urn:lsid:ipni.org:names:77204085-1(7) Pseudopodospermumelatum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraelata Boiss., Diagn. Pl. Orient., ser. 1, 4: 25. 1844. urn:lsid:ipni.org:names:77204086-1(8) Pseudopodospermumgracile (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneragracilis Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 721. 1964. urn:lsid:ipni.org:names:77204088-1(9) Pseudopodospermumhispanicum (L.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerahispanica L., Sp. Pl. 2: 791. 1753. urn:lsid:ipni.org:names:77204090-1(10) Pseudopodospermumhissaricum (C.Winkl.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerahissarica C.Winkl., Trudy Imp. S.-Peterburgsk. Bot. Sada 11: 172. 1889. urn:lsid:ipni.org:names:77204092-1(11) Pseudopodospermumidaeum (Gand.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Podospermumidaeum Gand., Bull. Soc. Bot. France 62: 155. 1916. urn:lsid:ipni.org:names:77204093-1(12) Pseudopodospermuminaequiscapum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerainaequiscapa Boiss., Fl. Orient. 3: 762. 1875. urn:lsid:ipni.org:names:77204091-1(13) Pseudopodospermumincisum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraincisa DC., Prodr. 7(1): 119. 1838. urn:lsid:ipni.org:names:77204089-1(14) Pseudopodospermuminconspicuum (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerainconspicua Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol., n.s. 42: 128. 1933. urn:lsid:ipni.org:names:77204087-1(15) Pseudopodospermumlamellatum (Krasch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneralamellata Krasch., Trudy Bot. Inst. Akad. Nauk S.S.S.R., ser. 1, Fl. Sist. Vyssh. Rast. 1: 180. 1933. urn:lsid:ipni.org:names:77204083-1(16) Pseudopodospermumleptophyllum (DC.) Kuth.(17) Pseudopodospermumlibanoticum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneralibanotica Boiss., Diagn. Pl. Orient., ser. 1, 11: 43. 1849. urn:lsid:ipni.org:names:77204081-1(18) Pseudopodospermummolle (M.Bieb.) Kuth.(19) Pseudopodospermummucidum  Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneramucida Rech.f., Aellen & Esfand., Oesterr. Bot. Z. 97: 264. 1950. urn:lsid:ipni.org:names:77204077-1(20) Pseudopodospermumovatum (Trautv.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraovata Trautv., Trudy Imp. S.-Peterburgsk. Bot. Sada 1: 275. 1871. urn:lsid:ipni.org:names:77204075-1(21) Pseudopodospermumpachycephalum (Podlech & Rech.f.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapachycephala Podlech & Rech.f. in Rechinger, Fl. Iranica 122: 36. 1977. urn:lsid:ipni.org:names:77204073-1(22) Pseudopodospermumpapposum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapapposa DC., Prodr. 7(1): 119. 1838. urn:lsid:ipni.org:names:77204072-1(23) Pseudopodospermumphaeopappum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Podospermumphaeopappum Boiss., Diagn. Pl. Orient., ser. 1, 7: 5. 1846. urn:lsid:ipni.org:names:77204070-1(24) Pseudopodospermumpubescens (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapubescens DC., Prodr. 7(1): 122. 1838. urn:lsid:ipni.org:names:77204067-1(25) Pseudopodospermumraddeanum (C.Winkl.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraraddeana C.Winkl., Trudy Imp. S.-Peterburgsk. Bot. Sada 11: 150. 1889. urn:lsid:ipni.org:names:77204062-1(26) Pseudopodospermumreverchonii (O.Debeaux ex Hervier) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerareverchonii O.Debeaux ex Hervier, Bull. Acad. Int. Géogr. Bot., sér. 3, 15(187–188): 107. 1905. urn:lsid:ipni.org:names:77204056-1(27) Pseudopodospermumsemicanum (DC.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerasemicana DC., Prodr. 7(1): 119. 1838. urn:lsid:ipni.org:names:77204055-1(28) Pseudopodospermumstrictum (Hornem.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerastricta Hornem., Hort. Bot. Hafn.: 2: 750. 1815. urn:lsid:ipni.org:names:77204052-1(29) Pseudopodospermumsuberosum (K.Koch) Kuth., Kavkaz. Predstav. Scorzonerinae: 94. 1978(30) Pseudopodospermumsyriacum (Boiss. & C.I.Blanche) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerasyriaca Boiss. & C.I.Blanche, Diagn. Pl. Orient., ser. 2, 3: 93. 1856. urn:lsid:ipni.org:names:77204049-1(31) Pseudopodospermumszowitzii (DC.) Kuth., Kavkaz. Predstav. Scorzonerinae: 92. 1978.(32) Pseudopodospermumtroodeum (Boiss.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneratroodea Boiss., Fl. Orient., suppl.: 320. 1888. urn:lsid:ipni.org:names:77204042-1(33) Pseudopodospermumturcomanicum (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraturcomanica Krasch. & Lipsch. in Lipschitz, Fragm. Monogr. Gen. Scorzonera 1: 80. 1935. urn:lsid:ipni.org:names:77204040-1(34) Pseudopodospermumundulatum (Vahl) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraundulata Vahl, Symb. Bot. 2: 86. 1791. urn:lsid:ipni.org:names:77204038-1(35) Pseudopodospermumviolaceum (Chamberlain) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraviolacea Chamberlain, Notes Roy. Bot. Gard. Edinburgh 33(2): 256. 1974. urn:lsid:ipni.org:names:77204037-1(36) N Africa: DZ; LY; MA; TN. Europe: AL; AT; BA; BE; BG; CH; CY; CZ; DE; ES; FR; GR; HR; HU; IT; MD; ME; MK; PL; PT; RO; RS; RU; SK; SI; UA. Asia-Temperate: AF; AM; AZ; EG; GE; IL; IQ; IR; JO; KW; KZ; LB; OM; SY; SA; TJ; TK; TR; UZ. Asia-Tropical: PK. Pseudopodospermumhispanicum as an edible crop was introduced in further regions.Pseudopodospermum, based on their morphological characters : Scorzoneraacuminata Boiss., S.aragatzii Kuth. ≡ Pseudopodospermumaragatzii (Kuth.) Kuth., S.coriacea A.Duran & Aksoy, S.crassifolia Krasch. & Lipsch., S.drarii Täckh., S.ferganica Krasch., S.gorovanica Nazarova, S.helodes Rech.f., S.karkasensis S.R.Safavi, S.lacera Boiss. & Balansa, S.limnophila Boiss., S.mariovoensis Micevski, S.mucida Rech.f. et al., S.pacis Güzel et al., S.nivalis Boiss. & Hausskn., S.rawii Rech.f. & Guest, S.scyria M.A.Gust. & Snogerup, S.serpentinica Rech.f., S.stenocephala Boiss., S.tadshikorum Krasch. & Lipsch., S.tunicata Rech.f. & Köie, S.tuzgoluensis A.Duran et al.The following species may belong to Taxon classificationPlantaeAsteralesAsteraceaeNazarova, Biol. Zhurn. Armenii 43: 179. 1990.7ADBE6F6-5D69-5409-BC6E-9468436F2166Scorzonerasect.Fibrillosae Nakai, Rep. Inst. Sci. Res. Manchoukuo, ser. 1, 6: 171. 1937. Type: Scorzoneraglabra Rupr. [= S.austriaca Willd.]  = Scorzonerasubsect.Egregia Kult., Tau-Sagyz Ekol. Osnovy Vvedeniya Ego Kul’t.: 108. 1938.  = Scorzonerasubg.Egregia (Kult.) Ovcz., Soobshch. Tadzhiksk. Fil. Akad. Nauk SSSR 20: 54. 1950.  ≡ Scorzonerasect.Egregia (Kult.) Lipsch., Fl. URSS 29: 58. 1964. Lectotype (designated here): Scorzoneratau-saghyz Lipsch. & G.G.Bosse  ≡ Scorzonerasect.Pusillae Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 59. 1939. Type: Scorzonerapusilla Pall.  = Scorzonerasect.Capito Tzvelev, Rast. Tsentral. Azii 14b: 115. 2009. Type: Scorzoneracapito Maxim.  = Takhtajanianthapusilla  NazarovaPerennials; caudex often with dark brown fibrous leaf sheath residues and base of the rosulate leaves often lanate; pollen with 6 lacunae.Habit, life form, subterranean parts: perennial herbs or subshrublets with a taproot or a tuber.Leaves: rosulate or crowded in the basal part of the stem, rarely dispersed along the stem, sessile or petiolate, filiform or ovate, apically sometimes hooked (T.pusilla), margin flat or undulate, entire or denticulate.Stem, synflorescence: stem solitary or branched, scape-like or leafy, capitula terminal.Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae, with 6 abporal lacunae  florets; florets yellow  with stout conglomerations; achene wall with parenchyma of two types, with thick- and thin-walled cells, present only as subepidermal continuous layer(s) or only with thick-walled cells, air cavities absent, sclerenchyma with continuous layer with a narrow or wider hunch on either side of the principal rib or, as in T.pseudodivaricata, without hunches, but discontinuous forming 5 bundles , or continuous, its fibres orientated parallel to the fruit axis, tannins absent.Pappus: 7–28 mm, ice white or dirty white, bristles scabrid in upper part.Chromosome number: x = 7 (diploids); 2n = 28 in T.pusilla (amphiploid); in T.tau-saghyz polyploid cytoraces 2n = 14, 21, 28, 42 and aneuploids with chromosome numbers 15, 16, 17, 18, 22 and 24, 28 and 42 are known (re known .Species.Takhtajanianthaaustriaca (Willd.) Zaika, Sukhor. & N.Kilian, comb. nov. : 1498. 1803 (incl. S.glabra Rupr.). urn:lsid:ipni.org:names:77204033-1(1) ov. Fig.  ≡ ScorzoTakhtajanianthacapito (Maxim.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracapito Maxim., Bull. Acad. Imp. Sci. Saint-Pétersb. 32: 491. 1888. urn:lsid:ipni.org:names:77204030-1(2) Takhtajanianthacrispa (M.Bieb.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneracrispa M.Bieb., Fl. Taur.-Caucas. 2: 234. 1808. urn:lsid:ipni.org:names:77204028-1(3) Takhtajanianthaikonnikovii (Krasch. & Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraikonnikovii Krasch. & Lipsch. in Lipschitz, Fragm. Monogr. Gen. Scorzonera 1: 109. 1935. urn:lsid:ipni.org:names:77204024-1(4) Takhtajanianthamongolica (Maxim.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneramongolica Maxim., Bull. Acad. Imp. Sci. Saint-Pétersb. 32: 492. 1888. urn:lsid:ipni.org:names:77204020-1(5) Takhtajanianthapseudodivaricata (Lipsch.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzonerapseudodivaricata Lipsch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 42: 158. 1933. urn:lsid:ipni.org:names:77204019-1(6) Takhtajanianthapusilla  Nazarova(7) Takhtajanianthasubacaulis (Regel) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraaustriacavar.subacaulis Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 6: 323. 1880. urn:lsid:ipni.org:names:77204015-1(8) Takhtajanianthatau-saghyz (Lipsch. & G.G.Bosse) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneratau-saghyz Lipsch. & G.G.Bosse, Trudy Nauchno-Issl. Lab. Tresta "Kauchukonos" 4: 27. 1930. urn:lsid:ipni.org:names:77204010-1(9) Takhtajanianthaveresczaginii (Kamelin & S.V.Smirn.) Zaika, Sukhor. & N.Kilian, comb. nov. ≡ Scorzoneraveresczaginii Kamelin & S.V.Smirn., Turczaninowia 5(1): 17. 2002. urn:lsid:ipni.org:names:77204008-1(10) N Africa: EG. Asia-Temperate: AF; AR; AZ; CN; IL; IR; JO; KG; KP; KR; KZ; MN; RU (Asiatic part); SA; TJ; TM; UZ. Asia-Tropical: PK. Europe: AL; AT; BA; BG; BY; CH; CZ; DE; FR; GR; HR; HU; IT; MD; ME; RO; RS; RU; SI; SK; UA.Takhtajaniantha, based on their morphological characters :The following species may belong to Scorzoneraaniana N.Kilian, S.grubovii Lipsch., S.karataviensis Kult. (possibly a synonym of T.tau-saghyz), S.manshurica Nakai, S.pamirica C.Shih, S.sinensis (Lipsch. & Krasch.) NakaiTaxon classificationPlantaeAsteralesAsteraceaeL., Sp. Pl.: 790. 1753.209CDCFE-7223-51AF-9029-5928108FB2B4Scorzonerahumilis L.Green in : 177: ScScorzonera in this revised sense as a monophyletic genus includes four larger clades, the Scorzonera s.str. clade (3 spp.), Scorzoneraalbicaulis clade (~17 spp.), Podospermum clade (~20 spp.) and Scorzonerapurpurea clade (2 spp.). In addition, it includes a few, so far somewhat isolated species . The genus in this cricumscription still shows considerable variation. Our analysis is insufficient with respect to a revision of the elaborated infrageneric classifications of previous authors. We therefore refer here to the clades of our trees.Taxon classificationPlantaeAsteralesAsteraceaes.str. cladeCC342938-05E6-5C98-8DBC-E81888283F93Scorzonerasect.ParvifloraeScorzoneraser.Parviflorae Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 720. 1964 [S.subsect.Parviflora Lipsch., Fragm. Monogr. Gen. Scorzonera 2: 112. 1939, nom. inval. .Inner phyllaries apically often with dark red or blackish spot; achene sclerenchyma insular in principal ribs below sclerenchyma and in secondary ribs above sclerenchyma  or leafy, glabrous or arachnoid-hairy becoming almost glabrous at the fruiting; capitula terminal, solitary or several.Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae; with 6 abporal lacunae ; achene wall with parenchyma insular in principal ribs below sclerenchyma and in secondary ribs above sclerenchyma, sclerenchyma continuous, its fibres orientated parallel to the achene axis, air cavities and tannins absent.Pappus: 11–16 mm, dirty white or rarely ice-white (S.parviflora), bristles plumose or apically scabrid (or 5–10 longer bristles scabrid and other bristles plumose).x = 7, diploids.Scorzoneraaristata Ramond ex DC.(1) Scorzonerahumilis L.   L. Fig. Scorzoneraparviflora Jacq.(3) Asia-Temperate: AF; AM; AR; CN; CY; GE; IR; KG; LB; MN; RU (Asiatic part); SY; TM; TR; UZ. Europe: AT; BE; BY; CH; CZ; DE; DK; EE; ES; FI; FR; GB; HR; IT; LT; LV; NL; NO; PL; PT; RO; RS; RU; SE; SK; SI; UA.Scorzoneraradiata Fisch. ex Ledeb. may also belong to this clade on account of its resemblance in morphological and carpological characters.Taxon classificationPlantaeAsteralesAsteraceaeclade5F63FF32-413D-5312-9D16-1811DF1FCFA5Podospermum DC. in Lamarck & Candolle, Fl. Franç., ed. 3, 4: 61. 1805, nom. cons.  ≡ Scorzonerasect.Podospermum (DC.) Benth. in Bentham & Hooker, Gen. Pl. 2: 532. 1873.  ≡ Scorzonerasubg.Podospermum (DC.) Lipsch., Fragm. Monogr. Gen. Scorzonera 1: 7. 1935. Type: Podospermumlaciniatum (L.) DC.  ≡ Herbs, biennial or perennial; leaves (at least some) pinnately divided; phyllaries in several series, often subapically corniculate; achene with conspicuous cylindrical carpopodium usually 1/5–1/3 as long as achene body; achene surface mostly smooth, more rarely verrucose or undulate and glabrous or rarely somewhat hairy; pollen with 18 or 24 lacunae; sclerenchyma in achene mesophyll of diverse orientation .Habit, life form, subterranean parts: biennial or perennial herbs, with taproot, often with woody caudex.Leaves: glabrous or pubescent, mostly located in the lower third of the stem, rosulate leaves usually present. Heterophylly often present sessile, lobate to entire).Stem, synflorescence: stem solitary or branched, leafy , glabrous or pubescent, capitula terminal.Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae; with 24 , glabrous or with scattered pubescence; achene wall with parenchyma continuous, above sclerenchyma and also well-expressed (continuous or discontinuous) in the principal ribs below sclerenchyma), sclerenchyma continuous, diversely orientated , air cavities and tannins absent.Pappus: 5–14 mm, white or dirty white, bristles plumose and only apically scabrid.x = 7, diploids.Scorzoneraalpigena (K.Koch) Grossh.(1) Scorzoneraarmeniaca (Boiss. & A.Huet) Boiss.(2) Scorzoneracana (C.A.Mey.) Griseb.(3) Scorzoneragrossheimii Lipsch. & Vassilcz.(4) Scorzonerahieraciifolia Hayek(5) Scorzonerakandavanica Rech.f.(6) Scorzonerakirpicznikovii Lipsch.(7) Scorzoneralachnostegia (Woronow) Lipsch.(8) Scorzoneralaciniata L.(9) Scorzoneraluristanica Rech.f.(10) Scorzonerameshhedensis (Rech.f.) Rech.f.(11) Scorzonerameyeri (K.Koch) Lipsch.  ch. Fig. Scorzonerapersepolitana Boiss.(13) Scorzoneraradicosa Boiss.(14) Scorzoneraschischkinii Lipsch. & Vassilcz.(15) Scorzonerasongorica (Kar. & Kir.) Lipsch. & Vassilcz.(16) Africa: DZ; EG; ES (Canary Islands); LY; MA; TN. Asia-Temperate: AF; AM; AZ; CN; CY; GE; GR; IL; IQ; IR; JO; KG; KZ; LB; RU (Asiatic part); SY; TJ; TR; TM; UZ. Asia-Tropical: IN; PK. Europe: all countries.Scorzoneralaciniata is introduced in Australasia (AU); North America (US); South America (AR).Podospermum clade based on their morphological characters: Scorzoneragrigoraschvilii (Sosn.) Lipsch., S.idae (Sosn.) Lipsch., S.lipschitzii (Kuth.) Czerep.The following species may also belong to the Taxon classificationPlantaeAsteralesAsteraceaeclade850D3E8E-61F8-5682-AD30-BA07D25B7EBEAchyroseris Sch.Bip., Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 21: 165. 1845. Type: Achyroserismacrosperma (Turcz. ex DC.) Sch.Bip. . Note: Achyroseris from Scorzonera on the “paleate receptacle” in A.macrosperma. This is, however, clearly erroneous, because a paleate (bracteate) receptacle is not present in any member of the Scorzonerinae; at most, the receptacle can be slightly hairy in some members of the phylogenetically distant Gelasia.  = Scorzonerasect.Piptopogon C.A.Mey., Bull. Soc. Imp. Naturalistes Moscou 21(3): 97. 1848.  = Scorzonerasubg.Piptopogon (C.A.Mey.) C.Díaz & Blanca, Anales Jard. Bot. Madrid 43: 330. 1987. Type: Scorzoneramacrosperma Turcz. ex DC.   ≡ Scorzonerasect.Turkestanicae Lipsch. in Bobrov & Tzvelev, Fl. URSS 29: 720. 1964. Type: Scorzoneraturkestanica Franch.  = Perennial herbs and subshrubs; pollen with 24 lacunae; achenes beaked; pappus dirty yellow, caducous.Habit, life form, subterranean parts: perennial herbs and subshrubs with a taproot and often with caudex.Leaves: rosulate and cauline, numerous, usually sessile (rosulate leaves petiolate in S.franchetii), semi-amplexicaule, linear to narrowly oblong, more often lanceolate, entire or crisp, glabrous or slightly pubescent.Stem, synflorescence: stem solitary or several, usually leafy but bracteate in S.acanthoclada and S.racemosa, capitula terminal and solitary or numerous, in spiciform or corymbiform synflorescence.Pollen: echinolophate, tricolporate and each colpus divided into 2 lacunae; with 24  lacunae , 1.5–2 times exceeding the involucre.Achenes: 7–45 mm, straight, without carpopodium, with more or less expressed beak, 10- or rarely 5-ribbed, papillate; achene wall with parenchyma well-expressed and represented by collenchyma-like cells, then present only as subepidermal continuous layers or sometimes parenchyma absent or discontinuous in the rib areas, insular in principal ribs below sclerenchyma and in secondary ribs above sclerenchyma or absent, air cavities absent, sclerenchyma present as layers with a gap in the principal ribs or continuous sclerenchymatous layers, its fibres orientated parallel to the fruit axis, tannins absent.Pappus: 6–28 mm, yellowish, bristles plumose below and scabrid in upper part.x = 7, diploids.Scorzoneraacanthoclada Franch.(1) Scorzoneraalbicaulis Bunge(2) Scorzonerabaldschuanica Lipsch.(3) Scorzonerabracteosa C.Winkl.(4) Scorzoneracrassicaulis Rech.f.(5) Scorzonerafranchetii Lipsch.(6) Scorzoneragraminifolia L.(7) Scorzonerapetrovii Lipsch.(8) Scorzoneraracemosa Franch.(9) Scorzoneratragopogonoides Regel & Schmalh.(10) Scorzoneratransiliensis Popov(11) Scorzoneraturkestanica Franch.(12) Scorzoneravirgata DC.(13) Africa: MA. Asia-Temperate: AF; RU (Asian part); CN; IR; KG; KP; KR; KZ; MN; TM; TJ; UZ. Asia-Tropical: IN; PK. Europe: ES; PT.Scorzoneraalaica Lipsch. may also belong to this clade based on morphological features.Taxon classificationPlantaeAsteralesAsteraceaecladeA3291AE4-CE61-5239-8369-AED7B5136CA9Scorzonerasect.PurpureaScorzonera 2: 104. 1939. Type  lacunae  ribs or flattened-triangular, glabrous or papillate: achene wall with parenchyma scarcely represented by one or several layers above and below sclerenchyma, of two types , air cavities absent, sclerenchyma continuous, of equal thickness throughout, sclerenchymatous fibres orientated parallel to the achene axis, tannins absent.Pappus: 7–13 mm, dirty white, bristles plumose in lower part and apically scabrid, 5–10 of them clearly longer than other bristles.x = 7, diploids.Scorzonerapurpurea L.(1) Scorzonerarosea Waldst. & Kit.(2) Scorzonerarhodantha Hausskn.(3) Asia-Temperate: KZ; RU (South Siberia). Asia-Tropical: IN; PK. Europe: AL; AT; BA; BG; CZ; DE; FR; GR; HR; HU; IT; MD; ME; MK; PL; RO; RS; RU; SI; SK; UA.Scorzonerarenzii  is sister to the Scorzonerapurpurea clade in our nrITS tree and forms a polytomy with the Scorzonerapurpurea and Podospermum clades in the plastid DNA tree. The spiciform synflorescences and other morphological features  rather indicate an affinity to the S.albicaulis clade.Scorzoneraangustifolia (pollen with 24 lacunae) and S.rupicola show some affinity to the S.albicaulis clade, but the second is resolved in the nrITS tree as sister to the polytomy including the S.purpurea and S.albicaulis clades, whereas the first is the third element of that polytomy.Scorzonerasect.Pentachlamys DC., Prodr. 7: 125. 1838, including two species from Nepal, S.bupleuroides D.Don, Prodr. Fl. Nepal.: 162. 1825 and S.roylei DC., Prodr. 7: 125. 1838, do not refer to Scorzonera or one of the genera segregated here. The holotype of S.roylei in G-DC (G00498633) is a species of Tragopogon. Original material of S.bupleuroides seems to be lost and the species is hardly a member of Scorzonera or one of its segregates as was already concluded by


Plasmodium falciparum nicotinamide mononucleotide adenylyltransferase: oligomeric assemblyStructural insights into ", DOI number: 10.1590/0074-02760180073, published in Mem Inst Oswaldo Cruz, Rio de Janeiro, Vol. 113(9): e180073, 2018, on page 1:In the article “Where it reads:ORIGINAL ARTICLEIt should read:SHORT COMMUNICATIONhttp://dx.doi.org/10.1590/0074-02760180073ER


The correct name is: Ramona M. Rodriguez. The correct citation is: McGaughey KD, Yilmaz-Swenson T, Elsayed NM, Cruz DA, Rodriguez RM, Kritzer MD, et al. (2018) Comparative evaluation of a new magnetic bead-based DNA extraction method from fecal samples for downstream next-generation 16S rRNA gene sequencing. PLoS ONE 13(8): e0202858.


The correct initials are Lima MG. The correct citation is: Lima MG, Malta DC, Monteiro CN, da Silva Sousa NF, Stopa SR, Medina LdPB, et al. (2019) Leisure-time physical activity and sports in the Brazilian population: A social disparity analysis. PLoS ONE 14(12): e0225940.


Expanding the single-visit approach for cervical cancer prevention: successes and lessons from Burkina Faso. Glob Health Sci Pract. 2018;6(2):288–298. https://doi.org/10.9745/GHSP-D-17-00328Harvey SA. Observe before you leap: why observation provides critical insights for formative research and intervention design that you'll never get from focus groups, Interviews, or KAP Surveys. Glob Health Sci Pract. 2018;6(2):299–316. https://doi.org/10.9745/GHSP-D-17-00471. (French translation of the abstract added)Koffi TB, Weidert K, Ouro Bitasse E, et al. Engaging men in family planning: perspectives from married men in Lomé, Togo. Glob Health Sci Pract. 2018;6(2):317–329. https://doi.org/10.9745/GHSP-D-17-00440Subramanian L, Simon C, Daniel EE. Increasing contraceptive use among young married couples in Bihar, India: evidence from a decade of implementation of the PRACHAR Project. Glob Health Sci Pract. 2018;6(2):330–344. https://doi.org/10.9745/GHSP-D-16-00290Sarma S, Nemser B, Cole-Lewis H, et al. Effectiveness of SMS technology on timely community health worker follow-up for childhood malnutrition: a retrospective cohort study in sub-Saharan Africa. Glob Health Sci Pract. 2018;6(2):345–355. https://doi.org/10.9745/GHSP-D-17-00427Marks KJ, Luthringer CL, Ruth LJ, et al. Review of grain fortification legislation, standards, and monitoring documents. Glob Health Sci Pract. 2018;6(2):356–371. https://doi.org/10.9745/GHSP-D-17-00417. (French translation of the abstract added)Ndiaye K, Portillo E, Ouedraogo D, Mobley A, Babalola S. High-risk advanced maternal age and high parity pregnancy: tackling a neglected need through formative research and action. Glob Health Sci Pract. 2018;6(2):372–383. https://doi.org/10.9745/GHSP-D-17-00318Kheang T, Lin MA, Lwin S, et al. Malaria case detection among mobile populations and migrant workers in Myanmar: comparison of 3 service delivery approaches. Glob Health Sci Pract. 2018;6(2):384–389. https://doi.org/10.9745/GHSP-D-18-00012Choi Y, Short Fabic M. Monitoring progress in equality for the Sustainable Development Goals: a case study of meeting demand for family planning. Glob Health Sci Pract. 2018;6(2):390–401. We have added French translations of the abstracts for a number of articles from the June 2018 issue  in which the content focused on countries where the official language is French. This has affected the page numbering of these and subsequent articles. The new citations are as follows:


N-acetyltransferase (SpeG) acetylates and thus neutralizes toxic polyamines. Studies indicate that SpeG plays an important role in virulence and pathogenicity of many bacteria, which have evolved SpeG-dependent strategies to control polyamine concentrations and survive in their hosts. In Escherichia coli, the two-component response regulator RcsB is reported to be subject to Nε-acetylation on several lysine residues, resulting in reduced DNA binding affinity and reduced transcription of the small RNA rprA; however, the physiological acetylation mechanism responsible for this behavior has not been fully determined. Here, we performed an acetyltransferase screen and found that SpeG inhibits rprA promoter activity in an acetylation-independent manner. Surface plasmon resonance analysis revealed that SpeG can physically interact with the DNA-binding carboxyl domain of RcsB. We hypothesize that SpeG interacts with the DNA-binding domain of RcsB and that this interaction might be responsible for SpeG-dependent inhibition of RcsB-dependent rprA transcription. This work provides a model for SpeG as a modulator of E. coli transcription through its ability to interact with the transcription factor RcsB. This is the first study to provide evidence that an enzyme involved in polyamine metabolism can influence the function of the global regulator RcsB, which integrates information concerning envelope stresses and central metabolic status to regulate diverse behaviors.Spermidine  N-acetyltransferase (GNAT) family, is a bacterial spermidine N-acetyltransferase that acetylates spermidine and spermine. These polyamines are toxic to bacteria at high concentrations and acetylation neutralizes this toxicity . A mtor RcsB . The inntor RcsB –24. RcsBtor RcsB . Both metor RcsB , 25–28.rprA , RcsB from Erwinia amylovora (RcsB-Er_am) [GI:33357861], YjjQ from E. coli (YjjQ-Es_co) [GI:83288197], BglJ from E. coli (BglJ-Es_co) [GI:3915634], YahA from E. coli (YahA-Es_co) [GI:2506596], YuaB from E. coli (YuaB-Es_co) [GI:81783897], DctR from E. coli (DctR-Es_co) [GI:57012697], RcsA from E. coli (RcsA-Es_co) [GI:60393000], EntR from Citrobacter freundii (EntR-Ci_fr) [GI:6015049], FimW from Salmonella enterica (FimW-Sa_en) [GI:585140], LuxR from Bacteroides thetaiotoamicron (LuxR-Ba_th) [GI:171849138], YgeK from E. coli (YgeK-Es_co) [GI:20140955], UhpA from E. coli (UhpA-Es_co) [GI:84029412], UvrY from E. coli (UvrY-Es_co) [GI:83288180], PA0034 from Pseudomonas aeruginosa (PA0034-Ps_ae) [GI:13959718], BvgA from Bordetella pertussis (BvgA-Bo_pe) [GI:61219948], FimZ from E. coli (FimZ-Es_co) [GI:84028128], EvgA from E. coli (EvgA-Es_co) [GI:82581667], FixJ from Sinorhizobium meliloti (FixJ-Si_me) [GI:159163516], StyR from P. fluorescens (StyR-Ps_fl) [GI:78100993], NodW from Bradyrhizobium diazoefficiens (NodW-Br_di) [GI:128495], Ycf29 from Porphyra purpurea (Ycf29-Po_pu) [GI:1723332], Ycf29 from Cyanophora paradoxa (Ycf29-Cy_pa) [GI:1351750], NarL from E. coli (NarL-Es_co) [GI:24158735], NarP from E. coli (NarP-Es_co) [GI:400374], GerE from Bacillus subtilis (GerE-Ba_su) [GI:13786948], VraR from Staphylococcus aureus (VraR-St_au) [GI:166007196], LiaR from B. subtilis (LiaR-Ba_su) [GI:68051995], DegU from B. subtilis (DegU-Ba_su) [GI:118438], YxjL from B. subtilis (YxjL-Ba_su) [GI:20141933], YhjB from E. coli (YhjB-Es_co) [GI:586682], CsgD from E. coli (CsgD-Es_co) [GI:1706166], MoaR from Enterobacter aerogenes (MoaR-En_ae) [GI:1709068], MalT from E. coli  [GI:189028606], SgaR from Hyphomicrobium methylovorum (SgaR-Hy_me) [GI:6094276], Rv08090c from Mycobacterium tuberculosis (Rv08090c-My_tu) [GI:6137301], AgmR from P. aeruginosa (AgmR-Ps_ae) [GI:121420], AlkS from P. oleovorans (AlkS-Ps_ol) [GI:6226550], ComA from B. subtilis (ComA-Ba_su) [GI:116903], YdfI from B. subtilis (YdfI-Ba_su) [GI:68566110], ExeN from Aeromonas salmonicida (ExeN-Ae_sa) [GI:1175862], LuxR from Aliivibrio fischeri (LuxR-Al_fi) [GI:462556], VanR from Vibrio anguillarum (VanR-Vi_an) [GI:9297072], SolR from Ralstonia solanacearum (SolR-Ra_so) [GI:9297032], AhyR from Aeromonas hydrophila (AhyR-Ae_hy) [GI:61218504], LasR from P. aeruginosa (LasR-Ps_ae) [GI:125980], Y4HQ from Sinorhizobium fredii (Y4HQ-Si_fr) [GI:2495427], SdiA from E. coli (SdiA-Es_co) [GI:2506570], PhzR from P. fluorescens (PhzR-Ps_fl) [GI:2495423], CarR from Pectobacterium carotovorum (CarR-Pe_ca) [GI:2495418], YenR from Yersinia enterocolitica (YenR-Ye_en) [GI:1723596], RhiR from Rhizobium leguminosarum (RhiR-Rh_le) [GI:417645], TraR from S. fredii (TraR-Si_fr) [GI:158429605], MoxX from Paracoccus denitrifican (MoxX-Pa_de) [GI:266552], BrpA from Streptomyces hygroscopicus (BrpA-St_hy) [GI:231653], RaiR from Rhizobium etli (RaiR-Rh_et) [GI:9297035], TraR from Agrobacterium tumefaciens (TraR-Ag_tu) GI:23200109 and TraJ from E. coli (TraJ-Es_co) [GI:464931].Phylogenetic tree was created in ClustalW2 server ((PDF)Click here for additional data file.S4 FigThe SPR sensograms of SpeG and transcription regulator RcsA. The RcsA protein was injected in four dilution series. Duplicate measurements for each concentration indicated above SPR sensograms were performed.(PDF)Click here for additional data file.