This is a PDF file of an unedited manuscript that has been accepted for publication. As a service... more This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. Ragweeds and relatives: molecular phylogenetics of Ambrosiinae (Asteraceae).
Botanical Journal of the Linnean Society, Jun 18, 2019
Stachydeae, comprising c. 470 species, are one of the most diverse and taxonomically puzzling gro... more Stachydeae, comprising c. 470 species, are one of the most diverse and taxonomically puzzling groups in Lamioideae. In the present study, the phylogenetic relationships in the Eurystachys clade (a phylogenetic name for all genera attributed to Stachydeae except Melittis) were reconstructed utilizing nuclear ribosomal DNA sequences (nrETS, 5S-NTS) from 148 accessions in 12 genera. Our phylogenetic results recovered Stachys as paraphyletic with numerous traditionally recognized genera nested in it. A broadly defined Eurystachys clade, however, was monophyletic. Unlike previous studies, the present study was able to resolve the group into 12 well-supported clades, named here as (1) Eriostomum, (2) Stachys, (3) Prasium, (4) Setifolia, (5) Distantes, (6) Burgsdorfia, (7) Hesiodia, (8) Empedoclia, (9) Sideritis, (10) Marrubiastrum, (11) Swainsoniana and (12) Olisia. These 12 clades were formally named in a phylogenetic nomenclature for the Eurystachys clade. Several infrageneric units were retrieved as monophyletic, namely Sideritis sections Burgsdorfia, Empedoclia and Hesiodia, Sideritis subgenus Marrubiastrum and Stachys sections Eriostomum (including Stachys section Mucronata) and Setifolia. The findings of this study also provide the basis for a future formal classification, with two options: (1) splitting of the Eurystachys clade into 12 monophyletic genera, all of them based on pre-existing genus names and redefined to encompass additional taxa, but without clear morphological apomorphies; or (2) lumping of all segregates into a broadly defined Stachys, including widely recognized and well-defined segregates such as Prasium and Sideritis.
Micromeria Benth. (Lamiaceae, Nepetoideae) is a very common genus in the Mediterranean region. To... more Micromeria Benth. (Lamiaceae, Nepetoideae) is a very common genus in the Mediterranean region. To test the monophyly of the genus and to elucidate its phylogenetic placement within subtribe Menthinae (Dumort) Endl. of tribe Mentheae Dumort we performed parsimony analysis of trnK intron sequence data of 51 accessions representing 15 genera of Nepetoideae and two genera of subfamily Ajugoideae. Tree topology reveals a well‐supported “core group” indicating four distinct lineages. The first one comprises three species of Satureja L. s.str., the second one includes taxa of the genus Clinopodium L. from both the Old and the New World, paraphyletic with respect to Monarda L. and two species of Micromeria section Pseudomelissa Benth. A third group contains all samples of the remaining Micromeria species. Within this monophyly, a western lineage including taxa from NW Africa, the Balearic, and the Canary Islands, is sister to an eastern lineage with species distributed from SE Asia to the western Mediterranean. In a further clade the genera Thymbra L., Thymus L., and Origanum L. are grouped together. Combined analysis using a reduced dataset of trnK/trnL‐F sequences increased support for the infrageneric resolution within Micromeria. Based on the phylogenetic reconstructions there is evidence that the genus as currently circumscribed is polyphyletic. Results are discussed in the context of morphology, karyology, and biogeography, outlining the necessity of removing section Pseudomelissa from Micromeria.
Ononis L. (Fabaceae, Papilionoideae) is a common genus in the circum-Mediterranean region. To inf... more Ononis L. (Fabaceae, Papilionoideae) is a common genus in the circum-Mediterranean region. To infer phylogenetic relationships, 69 Ononis species were analyzed using plastid trnL-F and nuclear ITS DNA sequences. Trees resulting from maximum parsimony analysis and Bayesian inference provide evidence that Ononis is monophyletic but contradict the traditional subgeneric division in O. sects. Ononis and Natrix. However, many of the 22 subsections of the genus are well supported by the molecular data. Phylogenetic reconstructions indicate five major lineages within the genus, which are morphologically supported by peduncle length and flower color, characters which also confirm the polyphyly suggested for O. subsect. Reclinatae. Ononis subsects. Antiquae and Rhodanthae form a basally branching monophylum, confirming the findings of previous studies. An examination of the life strategies dominant in different clades and the climatic conditions in the habitats of the species in the light of the molecular data suggests that O. subsects. Ononis, Mauritanicae and Canariensis are secondarily perennial, with the latter two serving as examples of high altitude woodiness and insular woodiness.
The Journal of Allergy and Clinical Immunology, 1996
Traditional particle counting has been used to establish aeroallergen concentration. To follow th... more Traditional particle counting has been used to establish aeroallergen concentration. To follow the changes in allergen concentration during one 24 hour period the following studies were undertaken. Samples were collected every 3 hours over a 24 hour period on top of a 5-story building using a spinning concentrator (SpinCon). The SpinCon concentrated the contents of laM 3 of air into a 10ml sample. RW allergen was evaluated using a double polyclonal assay using NIH rabbit antisera and a ragweed allergen standard. Sample Pollen Count Allergen Cone. (grains/M 3) (ktg/ml) Noon 29 0
Recent phylogenetic analyses based on single gene and combined data sets have substantially incre... more Recent phylogenetic analyses based on single gene and combined data sets have substantially increased our knowledge of the phylogeny of Caryophyllales s.l., indicating that additional carnivorous families are related to this alliance. In earlier contributions towards a reassessment of inter- and infrafamilial relationships slowly evolving genes had been preferred for phylogenetic inference. The resulting tree topologies based on rbcL and 18S rDNA, however, were characterized by limited resolution, low internal support and topological incongruence. Therefore genomic regions evolving more rapidly have been used in subsequent studies. Comparative sequencing of the matK gene and the flanking trnK intron region as well as combined analyses based on plastid matK, atpB, rbcL, and nuclear 18S rDNA have effectively improved resolution and internal support. Tree topologies revealed Caryophyllales s.l. as monophyletic group and indicated a clear division into two sister clades, the "core" and the "non-core" Caryophyllales (with Rhabdodendraceae and Simmondsiaceae with unclear affinities). Contrary to the "core" group (with Asteropeiaceae and Physenaceae as successive sister groups), which corresponds largely to the previous circumscription of the order, the monophyly of "non-core" Caryophyllales comprising Polygonaceae, Plumbaginaceae, Frankeniaceae, and Tamaricaceae along with the carnivorous families Droseraceae, Nepenthaceae, Drosophyllaceae, Dioncophyllaceae, and Ancistrocladaceae are a recent discovery. Based on reliable tree topologies it is hypothesized that pitfall traps of Nepenthes and snap traps typical for Aldrovanda and Dionaea were derived from a common ancestor with adhesive flypaper traps. With exception of Triphyophyllum carnivory was secondarily lost in the remaining Dioncophyllaceae (Dioncophyllum, Habropetalum) and all taxa of Ancistrocladaceae.
Traditional Chinese Medicine (TCM), with its long history, is deeply rooted in the Chinese cultur... more Traditional Chinese Medicine (TCM), with its long history, is deeply rooted in the Chinese culture and represents one of the oldest forms of medical therapy in the world. TCM has been used for thousands of years in China for health maintenance, disease prevention, and used to a lesser extent for the application of a variety of clinical therapies. Today, the global use of TCM is rapidly increasing and over 130 countries in the world are using Chinese Herbal Medicine (Hsiao 2007). The majority of drugs harvested and processed in China are of plant origin. A highly developed industry delivers medical plant preparations to the local Chinese customers and exports plant-based drugs to Asia and developing markets in Europe, Canada, and the USA.
To investigate phylogenetic and biogeographic relationships all species of Digitalis and Isoplexi... more To investigate phylogenetic and biogeographic relationships all species of Digitalis and Isoplexis and one species of the outgroup genera Antirrhinum and Globularia each were analyzed using nuclear ITS and plastid trnL-F sequences. Phylogenetic trees resulting from separate analyses were highly congruent. Combined analysis revealed two major lineages, which mark an early split in the genus Digitalis. While sections Digitalis, Frutescentes and Globiflorae appear monophyletic, sect. Tubiflorae is polyphyletic and sect. Macranthae should be expanded due to paraphyly. Our results provide evidence that all species of the genus Isoplexis have a common origin and are embedded in Digitalis. Isoplexis therefore should be reduced to sectional rank. The phylogenetic placement combined with ecomorphological characters indicates that Isoplexis may be a bird-pollinated Tertiary relict. Results are discussed in the context of biogeography, chemotaxonomy and morphology.
FIGURE 12. Flower photographs of Genlisea subgenus Tayloria in situ, corolla shown in ventral and... more FIGURE 12. Flower photographs of Genlisea subgenus Tayloria in situ, corolla shown in ventral and lateral view. A, B, G. metallica, Itacambira. C, D, G. oligophylla, Serra do Cipó. E, F, G. uncinata, Serra do Sincorá. G, H, G. flexuosa, Itacambira. J, K, G. exhibitionista, Cachoeira da Fumaça. L, M, G. nebulicola, Casca D'Anta. N, O, G. lobata, cultivated plant. P, Q, G. violacea, Diamantina. R, S, G. violacea, Serra do Cipó. T, U, G. violacea, Ibitipoca. Scale bars 1 cm.
FIGURE 4. Genlisea metallica (all from P.M. Gonella et al. 293). A, habit. B, section of the scap... more FIGURE 4. Genlisea metallica (all from P.M. Gonella et al. 293). A, habit. B, section of the scape. C, section of the pedicel. D, bract and bracteoles. E, insertion of bract and bracteoles to the peduncle. F, calyx. G, corolla. H, seeds. Drawing by A. Fleischmann.
FIGURE 10. Genlisea lobata (all from F. Rivadavia et al. 517). A, habit. B, section of the scape.... more FIGURE 10. Genlisea lobata (all from F. Rivadavia et al. 517). A, habit. B, section of the scape. C, bract (centre) and bracteoles. D, insertion of bract and bracteoles to the peduncle. E, calyx. F, corolla. G, valve of the seed capsule. H, seeds. Drawing by A. Fleischmann.
FIGURE 9. Genlisea nebulicola (all from F. Rivadavia & M. Peixoto 864). A, habit. B, leaf. C, sec... more FIGURE 9. Genlisea nebulicola (all from F. Rivadavia & M. Peixoto 864). A, habit. B, leaf. C, section of the pedicel. D, bract (centre) and bracteoles. E, insertion of bract and bracteoles to the peduncle. F, calyx and entire pedicel. G, corolla. H, seeds. Drawing by A. Fleischmann.
FIGURE 7. Genlisea flexuosa (all from P.M. Gonella et al. 292). A, habit. B, section of the scape... more FIGURE 7. Genlisea flexuosa (all from P.M. Gonella et al. 292). A, habit. B, section of the scape. C, bract and bracteoles. D, corolla. E, mature infructescence. F, seeds. Drawing by A. Fleischmann.
The figwort genus Scrophularia (Scrophulariaceae), widespread across the temperate zone of the No... more The figwort genus Scrophularia (Scrophulariaceae), widespread across the temperate zone of the Northern Hemisphere, comprises about 250 species and is a taxonomically challenging lineage displaying large morphological and chromosomal diversity. Scrophularia has never been examined in a large-scale phylogenetic and biogeographic context and represents a useful model for studying evolutionary history in the context of reticulation. A comprehensively sampled phylogeny of Scrophularia was constructed, based on nuclear ribosomal (ITS) and plastid DNA sequences (trnQ-rps16 intergenic spacer, trnL-trnF region) of 147 species, using Bayesian inference and maximum likelihood approaches. Selected individuals were cloned. A combination of coding plastid indels and ITS intra-individual site polymorphisms, and applying Neighbor-Net and consensus network methods for adequate examination of within-dataset uncertainty as well as among-dataset incongruence, was used to disentangle phylogenetic relationships. Furthermore, divergence time estimation and ancestral area reconstruction were performed to infer the biogeographic history of the genus. The analyses reveal significant plastid-nuclear marker incongruence and considerable amounts of intra-individual nucleotide polymorphism in the ITS dataset. This is due to a combination of processes including reticulation and incomplete lineage sorting, possibly complicated by inter-array heterogeneity and pseudogenization in ITS in the presence of incomplete concerted evolution. Divergence time estimates indicate that Scrophularia originated during the Miocene in Southwestern Asia, its primary center of diversity. From there, the genus spread to Eastern Asia, the New World, Europe, Northern Africa, and other regions. Hybridization and polyploidy played a key role in the diversification history of Scrophularia, which was shaped by allopatric speciation in mountainous habitats during different climatic periods.
This is a PDF file of an unedited manuscript that has been accepted for publication. As a service... more This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. Ragweeds and relatives: molecular phylogenetics of Ambrosiinae (Asteraceae).
Botanical Journal of the Linnean Society, Jun 18, 2019
Stachydeae, comprising c. 470 species, are one of the most diverse and taxonomically puzzling gro... more Stachydeae, comprising c. 470 species, are one of the most diverse and taxonomically puzzling groups in Lamioideae. In the present study, the phylogenetic relationships in the Eurystachys clade (a phylogenetic name for all genera attributed to Stachydeae except Melittis) were reconstructed utilizing nuclear ribosomal DNA sequences (nrETS, 5S-NTS) from 148 accessions in 12 genera. Our phylogenetic results recovered Stachys as paraphyletic with numerous traditionally recognized genera nested in it. A broadly defined Eurystachys clade, however, was monophyletic. Unlike previous studies, the present study was able to resolve the group into 12 well-supported clades, named here as (1) Eriostomum, (2) Stachys, (3) Prasium, (4) Setifolia, (5) Distantes, (6) Burgsdorfia, (7) Hesiodia, (8) Empedoclia, (9) Sideritis, (10) Marrubiastrum, (11) Swainsoniana and (12) Olisia. These 12 clades were formally named in a phylogenetic nomenclature for the Eurystachys clade. Several infrageneric units were retrieved as monophyletic, namely Sideritis sections Burgsdorfia, Empedoclia and Hesiodia, Sideritis subgenus Marrubiastrum and Stachys sections Eriostomum (including Stachys section Mucronata) and Setifolia. The findings of this study also provide the basis for a future formal classification, with two options: (1) splitting of the Eurystachys clade into 12 monophyletic genera, all of them based on pre-existing genus names and redefined to encompass additional taxa, but without clear morphological apomorphies; or (2) lumping of all segregates into a broadly defined Stachys, including widely recognized and well-defined segregates such as Prasium and Sideritis.
Micromeria Benth. (Lamiaceae, Nepetoideae) is a very common genus in the Mediterranean region. To... more Micromeria Benth. (Lamiaceae, Nepetoideae) is a very common genus in the Mediterranean region. To test the monophyly of the genus and to elucidate its phylogenetic placement within subtribe Menthinae (Dumort) Endl. of tribe Mentheae Dumort we performed parsimony analysis of trnK intron sequence data of 51 accessions representing 15 genera of Nepetoideae and two genera of subfamily Ajugoideae. Tree topology reveals a well‐supported “core group” indicating four distinct lineages. The first one comprises three species of Satureja L. s.str., the second one includes taxa of the genus Clinopodium L. from both the Old and the New World, paraphyletic with respect to Monarda L. and two species of Micromeria section Pseudomelissa Benth. A third group contains all samples of the remaining Micromeria species. Within this monophyly, a western lineage including taxa from NW Africa, the Balearic, and the Canary Islands, is sister to an eastern lineage with species distributed from SE Asia to the western Mediterranean. In a further clade the genera Thymbra L., Thymus L., and Origanum L. are grouped together. Combined analysis using a reduced dataset of trnK/trnL‐F sequences increased support for the infrageneric resolution within Micromeria. Based on the phylogenetic reconstructions there is evidence that the genus as currently circumscribed is polyphyletic. Results are discussed in the context of morphology, karyology, and biogeography, outlining the necessity of removing section Pseudomelissa from Micromeria.
Ononis L. (Fabaceae, Papilionoideae) is a common genus in the circum-Mediterranean region. To inf... more Ononis L. (Fabaceae, Papilionoideae) is a common genus in the circum-Mediterranean region. To infer phylogenetic relationships, 69 Ononis species were analyzed using plastid trnL-F and nuclear ITS DNA sequences. Trees resulting from maximum parsimony analysis and Bayesian inference provide evidence that Ononis is monophyletic but contradict the traditional subgeneric division in O. sects. Ononis and Natrix. However, many of the 22 subsections of the genus are well supported by the molecular data. Phylogenetic reconstructions indicate five major lineages within the genus, which are morphologically supported by peduncle length and flower color, characters which also confirm the polyphyly suggested for O. subsect. Reclinatae. Ononis subsects. Antiquae and Rhodanthae form a basally branching monophylum, confirming the findings of previous studies. An examination of the life strategies dominant in different clades and the climatic conditions in the habitats of the species in the light of the molecular data suggests that O. subsects. Ononis, Mauritanicae and Canariensis are secondarily perennial, with the latter two serving as examples of high altitude woodiness and insular woodiness.
The Journal of Allergy and Clinical Immunology, 1996
Traditional particle counting has been used to establish aeroallergen concentration. To follow th... more Traditional particle counting has been used to establish aeroallergen concentration. To follow the changes in allergen concentration during one 24 hour period the following studies were undertaken. Samples were collected every 3 hours over a 24 hour period on top of a 5-story building using a spinning concentrator (SpinCon). The SpinCon concentrated the contents of laM 3 of air into a 10ml sample. RW allergen was evaluated using a double polyclonal assay using NIH rabbit antisera and a ragweed allergen standard. Sample Pollen Count Allergen Cone. (grains/M 3) (ktg/ml) Noon 29 0
Recent phylogenetic analyses based on single gene and combined data sets have substantially incre... more Recent phylogenetic analyses based on single gene and combined data sets have substantially increased our knowledge of the phylogeny of Caryophyllales s.l., indicating that additional carnivorous families are related to this alliance. In earlier contributions towards a reassessment of inter- and infrafamilial relationships slowly evolving genes had been preferred for phylogenetic inference. The resulting tree topologies based on rbcL and 18S rDNA, however, were characterized by limited resolution, low internal support and topological incongruence. Therefore genomic regions evolving more rapidly have been used in subsequent studies. Comparative sequencing of the matK gene and the flanking trnK intron region as well as combined analyses based on plastid matK, atpB, rbcL, and nuclear 18S rDNA have effectively improved resolution and internal support. Tree topologies revealed Caryophyllales s.l. as monophyletic group and indicated a clear division into two sister clades, the "core" and the "non-core" Caryophyllales (with Rhabdodendraceae and Simmondsiaceae with unclear affinities). Contrary to the "core" group (with Asteropeiaceae and Physenaceae as successive sister groups), which corresponds largely to the previous circumscription of the order, the monophyly of "non-core" Caryophyllales comprising Polygonaceae, Plumbaginaceae, Frankeniaceae, and Tamaricaceae along with the carnivorous families Droseraceae, Nepenthaceae, Drosophyllaceae, Dioncophyllaceae, and Ancistrocladaceae are a recent discovery. Based on reliable tree topologies it is hypothesized that pitfall traps of Nepenthes and snap traps typical for Aldrovanda and Dionaea were derived from a common ancestor with adhesive flypaper traps. With exception of Triphyophyllum carnivory was secondarily lost in the remaining Dioncophyllaceae (Dioncophyllum, Habropetalum) and all taxa of Ancistrocladaceae.
Traditional Chinese Medicine (TCM), with its long history, is deeply rooted in the Chinese cultur... more Traditional Chinese Medicine (TCM), with its long history, is deeply rooted in the Chinese culture and represents one of the oldest forms of medical therapy in the world. TCM has been used for thousands of years in China for health maintenance, disease prevention, and used to a lesser extent for the application of a variety of clinical therapies. Today, the global use of TCM is rapidly increasing and over 130 countries in the world are using Chinese Herbal Medicine (Hsiao 2007). The majority of drugs harvested and processed in China are of plant origin. A highly developed industry delivers medical plant preparations to the local Chinese customers and exports plant-based drugs to Asia and developing markets in Europe, Canada, and the USA.
To investigate phylogenetic and biogeographic relationships all species of Digitalis and Isoplexi... more To investigate phylogenetic and biogeographic relationships all species of Digitalis and Isoplexis and one species of the outgroup genera Antirrhinum and Globularia each were analyzed using nuclear ITS and plastid trnL-F sequences. Phylogenetic trees resulting from separate analyses were highly congruent. Combined analysis revealed two major lineages, which mark an early split in the genus Digitalis. While sections Digitalis, Frutescentes and Globiflorae appear monophyletic, sect. Tubiflorae is polyphyletic and sect. Macranthae should be expanded due to paraphyly. Our results provide evidence that all species of the genus Isoplexis have a common origin and are embedded in Digitalis. Isoplexis therefore should be reduced to sectional rank. The phylogenetic placement combined with ecomorphological characters indicates that Isoplexis may be a bird-pollinated Tertiary relict. Results are discussed in the context of biogeography, chemotaxonomy and morphology.
FIGURE 12. Flower photographs of Genlisea subgenus Tayloria in situ, corolla shown in ventral and... more FIGURE 12. Flower photographs of Genlisea subgenus Tayloria in situ, corolla shown in ventral and lateral view. A, B, G. metallica, Itacambira. C, D, G. oligophylla, Serra do Cipó. E, F, G. uncinata, Serra do Sincorá. G, H, G. flexuosa, Itacambira. J, K, G. exhibitionista, Cachoeira da Fumaça. L, M, G. nebulicola, Casca D'Anta. N, O, G. lobata, cultivated plant. P, Q, G. violacea, Diamantina. R, S, G. violacea, Serra do Cipó. T, U, G. violacea, Ibitipoca. Scale bars 1 cm.
FIGURE 4. Genlisea metallica (all from P.M. Gonella et al. 293). A, habit. B, section of the scap... more FIGURE 4. Genlisea metallica (all from P.M. Gonella et al. 293). A, habit. B, section of the scape. C, section of the pedicel. D, bract and bracteoles. E, insertion of bract and bracteoles to the peduncle. F, calyx. G, corolla. H, seeds. Drawing by A. Fleischmann.
FIGURE 10. Genlisea lobata (all from F. Rivadavia et al. 517). A, habit. B, section of the scape.... more FIGURE 10. Genlisea lobata (all from F. Rivadavia et al. 517). A, habit. B, section of the scape. C, bract (centre) and bracteoles. D, insertion of bract and bracteoles to the peduncle. E, calyx. F, corolla. G, valve of the seed capsule. H, seeds. Drawing by A. Fleischmann.
FIGURE 9. Genlisea nebulicola (all from F. Rivadavia & M. Peixoto 864). A, habit. B, leaf. C, sec... more FIGURE 9. Genlisea nebulicola (all from F. Rivadavia & M. Peixoto 864). A, habit. B, leaf. C, section of the pedicel. D, bract (centre) and bracteoles. E, insertion of bract and bracteoles to the peduncle. F, calyx and entire pedicel. G, corolla. H, seeds. Drawing by A. Fleischmann.
FIGURE 7. Genlisea flexuosa (all from P.M. Gonella et al. 292). A, habit. B, section of the scape... more FIGURE 7. Genlisea flexuosa (all from P.M. Gonella et al. 292). A, habit. B, section of the scape. C, bract and bracteoles. D, corolla. E, mature infructescence. F, seeds. Drawing by A. Fleischmann.
The figwort genus Scrophularia (Scrophulariaceae), widespread across the temperate zone of the No... more The figwort genus Scrophularia (Scrophulariaceae), widespread across the temperate zone of the Northern Hemisphere, comprises about 250 species and is a taxonomically challenging lineage displaying large morphological and chromosomal diversity. Scrophularia has never been examined in a large-scale phylogenetic and biogeographic context and represents a useful model for studying evolutionary history in the context of reticulation. A comprehensively sampled phylogeny of Scrophularia was constructed, based on nuclear ribosomal (ITS) and plastid DNA sequences (trnQ-rps16 intergenic spacer, trnL-trnF region) of 147 species, using Bayesian inference and maximum likelihood approaches. Selected individuals were cloned. A combination of coding plastid indels and ITS intra-individual site polymorphisms, and applying Neighbor-Net and consensus network methods for adequate examination of within-dataset uncertainty as well as among-dataset incongruence, was used to disentangle phylogenetic relationships. Furthermore, divergence time estimation and ancestral area reconstruction were performed to infer the biogeographic history of the genus. The analyses reveal significant plastid-nuclear marker incongruence and considerable amounts of intra-individual nucleotide polymorphism in the ITS dataset. This is due to a combination of processes including reticulation and incomplete lineage sorting, possibly complicated by inter-array heterogeneity and pseudogenization in ITS in the presence of incomplete concerted evolution. Divergence time estimates indicate that Scrophularia originated during the Miocene in Southwestern Asia, its primary center of diversity. From there, the genus spread to Eastern Asia, the New World, Europe, Northern Africa, and other regions. Hybridization and polyploidy played a key role in the diversification history of Scrophularia, which was shaped by allopatric speciation in mountainous habitats during different climatic periods.
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