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Maximum likelihood phylogram of Boraginaceae based on the trnL-trnF dataset, with Cordia (Cordiaceae), Myriopus and Tournefortia (Heliotropiaceae), and Codon and Wigandia (Hydrophyllaceae) as outgroups. Numbers above branches: Maximum likelihood bootstrap support values, numbers below branches: Bayesian posterior probabilities. Branch lengths are to scale. Species names in bold represent members of " Trigonotideae " . The four tribes (EC, Echiochileae) are shown in the broad sense (i.e. including the members of the former " Trigonotideae " ).
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Boraginaceae tribe Trigonotideae comprises a heterogenous assemblage of taxa, many of which have been shown to belong to widely divergent lineages in Boraginaceae in the recent past, with some taxa now assigned to three of the four currently recognized tribes of the Boraginaceae s.
s., namely the Cynoglosseae, Echiochileae, and Lithospermeae. The s...
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Context 1
... supported, contradicting tree topologies were not observed in single gene analyses, indi- cating parallel evolution of the two loci. The most compre- hensive sampling was used for the trnL-trnF analysis ( Fig. 4 ). Boraginaceae were monophyletic and fell into four well- supported clades: Echiochileae (100 ML bootstrap support value: LBS, 1.00 Bayesian posterior probability: BPP), Cyno- glosseae s. l. (100 LBS, 0.94 BPP), Boragineae (99 LBS, 0.96 BPP), and Lithospermeae (97 LBS, 0.99 BPP). ...
Context 2
... Prolithospermum also from Texas ( Thomasson 1977 ;Gabel et al. 1998 ). The nutlets of † Prolithospermum johnstonii are virtually indistinguishable in shape and size from those of western European Pentaglottis ( Thomasson 1979 ), and † Biorbia closely conforms to the morphology found, for example, in extant Pulmonaria (compare Gabel et al. 1998 , Fig. 4.6). The cicatrices of † Biorbia and † Prolithospermum are distinctly con- cave, indicating the presence of an elaiosome, lost during fos- silization, similar in shape and size to that of extant Eurasian Boragineae. All three fossil taxa share the strong develop- ment of the abaxial side of the nutlet with Moritzia and Thaumatocaryon ...
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10 taxa representing three groups of taxa in Iran, the N. caspic...
The taxonomic position and affinities of the rare Turkish endemic Arnebia purpurea are analyzed using nuclear and plastid DNA sequence data and morphological characters. Phylogenetic analysis of a wide sample of old-world Lithospermeae consistently retrieved a clade with this species sister to Huynhia pulchra, the only member of the genus Huynhia....
Palynological analysis of the genus Mertensia Roth, s. lat. (Boraginaceae) was performed. All pollen grains studied were mostly dumbbell shaped and had 6-heteroaperturate pollen type without exception. The type of pollen of Mertensia does not contradict recent classifications that assign Mertensia to Cynoglosseae. Among six sections of Mertensia, p...
Citations
... The basic fruit type is highly conserved, with the mericarpid always provided with a dry fruit wall enclosing the seed and thus technically representing nut fruits. However, the nut fruits formed by Boraginaceae show a tremendous range of specialized modifications and these have been central to the infrafamilial classification with individual groups characterized by particular modifications, such as the elaiosomes in Boragineae or the þ/− smooth and strongly mineralized pericarp of Lithospermeae (Weigend et al., 2009(Weigend et al., , 2010. Nutlet modifications are clearly geared towards different dispersal mechanisms, with epichory and anemochory particularly common, but instances of ant-dispersal and peculiar types of endochory have also been reported . ...
Recent phylogenetic studies have challenged the traditional classification of the subtribe Cynoglossinae, which was based on nutlet characteristics. To investigate the evolution of fruit traits related to dispersal modes in this complex group, we conducted a study using scanning electron microscopy to examine 28 taxa representing all previously recognized lineages of the subtribe. Cynoglossinae displays four main types of nutlets: marginate, emarginate, flat wing, and incurved wing. Our findings reveal the arrangement of glochids and their structure, including the number of apical hooks, and their surface ornamentation is highly variable both across and within these four main types. We reconstructed the phylogenetic tree with four main clades (i.e., Clades I–IV) using Maximum likelihood analysis. In addition, we mapped four nutlet traits (i.e., nutlet margin, nutlet glochid, glochid sculpture, and glochid hooks) associated with different dispersal modes using ancestral state reconstruction analysis. The ancestor of subtribe Cynoglossinae was inferred to have marginate and glochidiate nutlets, with glochids externally provided with complex spiny tubercles and bearing at least four hooks at the tip as well as epizoochory dispersal mode. These ancestral states have been retained in Clade II (East Asian-Australian-African Cynoglossum-Lindelofia clade) and Clade III (Mediterranean-Asian Cynoglossum-Solenanthus-Trachelanthus clade), with several independent shifts to emarginate fruits. Transitions from marginate to wide-winged nutlets have occurred at least four independent times, facilitating wind-assisted dispersal in Clade I (Microparacaryum clade), Clade IV (Paracaryum-Mattiastrum-Rindera clade), Rindera tetraspis, and Mattiastrum crista-galli (both belonging to Clade III), where a transition from tuberculate to papillate surface has also occurred.
... Finally, Weigend et al. (2010), in a study of tribal relationships in the Boraginoideae using trnL-F cpDNA sequences, obtained results similar to Långströ m and Chase (2002). Weigend et al. included two species of Cryptantha, two of Plagiobothrys, and one of Amsinckia. ...
... Given that the recent studies of Långströ m and Chase (2002) and Weigend et al. (2010) recognize an expanded tribe Cynoglosseae (inclusive of the tribe Cryptantheae of Schwarzer 2007), we recognize Cryptantha s. l. as belonging to subtribe Cryptanthinae (Brand 1931). We test whether this group corresponds to a clade comprised of Cryptantha s. l. and the genera Amsinckia, Pectocarya, and Plagiobothrys. ...
Cryptantha (Boraginaceae) is a group of approximately 200 annual and perennial species, representing two-thirds of the diversity within subtribe Cryptanthinae. The genus exhibits an amphitropic distribution, occurring in temperate and desert regions of western North and South America. Fifty samples of 45 species of Cryptantha s. l., exemplars of the related genera Amsinckia, Pectocarya, and Plagiobothrys, and four outgroup taxa were sequenced for two gene regions, the nuclear ribosomal gene, ITS, and the trnL UAA intron region of the chloroplast genome. These data were used to assess phylogenetic relationships using parsimony, maximum likelihood, and Bayesian inference methods. Cryptantha s. l. was found to be polyphyletic, with its members placed among several well-supported clades. Based on these analyses, we propose resurrection of the genera Eremocarya, Greeneocharis, Johnstonella, and Oreocarya, and recognition of a newly delimited Cryptantha s. s. The related genera Amsinckia and Pectocarya were resolved as monophyletic and most closely related to various clades within Cryptantha s. l. Plagiobothrys was resolved as polyphyletic in three clades, these clades corresponding to previously named sections or groups of sections. The Cryptanthinae is supported as monophyletic. Character trait analyses support the multiple, derived evolution of perenniality, reduction in nutlet number, nutlet heteromorphism, smooth nutlet sculpturing, heterostyly, and cleistogamy. Although sampling is incomplete, this study generally supports the hypothesis of repeated, unidirectional dispersal events, from North to South America. Genera resurrected include:
... However, molecular phylogenetic studies have not supported the status of the tribe Trigonotideae. Based on a few DNA markers such as atpB, trnL-trnF, matK, ndhF, and rbcL and nuclear ribosomal DNA internal transcribed spacer (nrITS) regions, previous phylogenetic analyses have indicated that Trigonotideae belongs to the tribe Cynoglosseae, and Trigonotis should be classified under Cynoglosseae (Långstr€ om and Chase, 2002;Weigend et al., 2010;Nazaire and Hufford, 2012). However, Weigend et al. (2013) and Chac on et al. (2016) revised Boraginaceae s.str. ...
... Detailed information about the samples used for nutlet morphology and chloroplast genome sequence analyses is provided in Tables S1 and S2, respectively. In addition, based on the phylogenetic results of Weigend et al. (2010Weigend et al. ( , 2013, Cohen (2014) and Chac on et al. (2016), we used the Lappula myosotis Moench (MZ_959108) as an outgroup to explore the phylogenetic relationships of Trigonotis. ...
... Furthermore, except for phylogenetic trees reconstructed based on IR and SSC regions, every node had strong support. The classifications of several Boraginaceae species and their evolutionary relationships has previously been inferred based on several DNA markers and morphological characters (Långstr€ om and Chase, 2002;Weigend et al., 2010Weigend et al., , 2013Cohen, 2014;Chac on et al., 2016). However, in previous phylogenetic studies, fewer Trigonotis taxa were sampled. ...
The genus Trigonotis comprises nearly 60 species mainly distributed in East and Southeast Asia. China has the largest number of Trigonotis species in the world, with a total of 44 species, of which 38 are endemic. Nutlet morphology is useful for the taxonomic delimitation of Trigonotis. However, there are still controversial circumscriptions of nutlet shape in some species. In previous studies, interspecies phylogenetic relationships were inferred using few DNA markers and very few taxa, which possibly led to erroneous or incomplete conclusions. In this study, the nutlet morphology of 39 Trigonotis taxa and the characteristics of 34 complete chloroplast genomes (29 taxa) were investigated and analyzed. Then, the phylogenetic relationships were discussed within this genus based on complete chloroplast genomes. To the best of our knowledge, this study is the first comprehensive analysis of nutlet morphology and complete chloroplast genome of Trigonotis. Based on nutlet morphology, Trigonotis can be divided into two groups: Group 1, hemispherical or oblique tetrahedron with carpopodiums, and Group 2, inverted tetrahedron without carpopodiums. The chloroplast genome of Trigonotis exhibited a typical quadripartite structure, including 84-86 protein-coding, 37 transfer RNA, and 8 ribosomal RNA genes, with a total length of 147,247-148,986 bp. Genes in the junctions were well conserved in Trigonotis, similar to those in other Boraginaceae s.str. species. Furthermore, Trigonotis chloroplast genomes showed relatively high diversity, with more conserved genic regions than intergenic regions; in addition, we detected 14 hot spots (Pi > 0.005) in non-coding regions. Phylogenetic analyses based on chloroplast genome data identified highly resolved relationships between Trigonotis species. Specifically, Trigonotis was divided into two clades with strong support: one clade included species with hemispherical or oblique tetrahedron nutlets with carpopodiums and bracts, whereas the other clade included species with inverted tetrahedron nutlets without carpopodiums or bracts. Our results may inform future taxonomic, phylogenetic, and evolutionary studies on Boraginaceae.
... For subsequent molecular dating analyses, the initial dataset of 70 accessions (Supplementary Fig. S3 1 ) was reduced to 56 accessions (one accession per species). Boraginaceae have an abundant fossil record (Weigend et al. 2010;Luebert et al. 2016), but fossils within Cynoglossinae have not yet been unearthed. Therefore, the divergence time was estimated based on a secondary calibration from a previous study by Chacón et al. (2017). ...
The Irano-Turanian (I-T) bioregion harbours one of the Old world’s greatest repositories of botanical diversity; however, the diversification patterns and the phenotypic evolution of its flora are sorely understudied. The subtribe Cynoglossinae is characteristic of the western I-T bioregion, species–rich both in the desertic lowlands and the more mesic highlands of the Iranian plateau. About 70 species of Cynoglossinae are present in the Iranian plateau, 47 of which are endemic to the plateau.Herein, nuclear ITS and cpDNA rpl32-trnL and trnH–psbA sequences were used to investigate the molecular phylogeny, historical biogeography and ancestral character states of Cynoglossinae. Molecular dating and ancestral range reconstruction analyses indicated that the subtribe Cynoglossinae has initiated its diversification from the eastern part of the western I-T during the mid-Miocene, concomitantly with the uplift of the Pamir and Hindu Kush mountains. Moreover, from the Pliocene onwards the Afghan-India collision and extensive deformation of the Arabia-Eurasia convergence probably promoted allopatric speciation in Cynoglossinae via mostly vicariance events. Evolution of annuals with small nutlets from perennials with large nutlets was accompanied by mesic to desert habitats shifts. Herein, to explain distribution of Cynoglossinae in the western I-T, the congruence between cladogenetic, geological and palaeoclimatic events was investigated.
... This family is characterized by a scorpioid cymose inflorescence (Buys and Hilger, 2003), a gynobasic style, and a two-part ovary that breaks into four nutlets. This circumscription is equivalent to, and has in the past been referred to as, Boraginaceae s.s. or Boraginoideae (Small, 1903;Diane et al. 2002;Weigend et al. 2010). This broader circumscription has included four taxa treated as either subfamilies (Boraginoideae, Cordioideae, Ehretioideae, and Heliotropioideae) or families that are characterized by a scorpioid cyme and two-parted gynoecium (style position and fruit type vary) (Lawrence, 1937;Cronquist, 1981;Al-Shehbaz, 1991;Takhtadzhian, 1997). ...
Heliotropium pakistanicum Shaheen sp. nov., has been described as new species to science. This taxon has been collected from Thal Desert, Punjab province, Pakistan. This species is closely associated with H. cabulicum Bunge. However, It differs from H. cabulicum in its leaf shape, inflorescence and glabrous corolla tube and bulbose base hairs. Taxonomic details of Heliotropium pakistanicum have been provided in this article.
... DNA extraction, amplification, and sequencing DNA extraction, amplification, purification, and sequencing followed standard protocols as described in Gottschling & Hilger (2001) and Weigend et al. (2010). Samples were sequenced for four genomic regions: the nuclear ribosomal ITS1-5.8S-ITS2 ...
Urtica L. (Urticaceae) is a subcosmopolitan genus, which is also common throughout Asia. Taxon differentiation in the Asian Urtica species is difficult due to the limited diversity of taxonomically useful characters combined with a range of phenotypic variation. The present study investigates the species Urtica thunbergiana Sieb. & Zucc. (incl. U. laetevirens) from eastern Asia based on herbarium collections, including most of the type specimens. The delimitation of U. thunbergiana is expanded to include all three currently recognized subspecies of U. laetevirens Maxim. Also, one newly infraspecific taxon is described from Taiwan. Based on our revision, the recognition of a total of four subspecies of Urtica thunbergiana is proposed: subsp. thunbergiana, subsp. dentata (Hand.-Mazz.) K.Becker & Weigend, subsp. silvatica (Hand.-Mazz.) K.Becker & Weigend, and subsp. perserrata, subspec. nov. The systematic re-arrangements are based on morphological analyses and a highly resolved phylogeny based on the molecular markers ITS1–5.8S–ITS2, psbA–trnH, trnL–trnF and trnS–trnG.
... DNA extraction, amplification, purification, and sequencing followed standard protocols as described in Gottschling & Hilger (2001) andWeigend et al. (2010). Samples were sequenced for four genomic regions: the nuclear ribosomal ITS1-5.8S-ITS2 ...
Urtica L. (Urticaceae) is a subcosmopolitan genus common throughout temperate Asia. Species delimitation is very difficult and the present study investigates a group of taxa associated with Urtica fissa from East-Southeast Asia and based mainly on herbarium collections, including most of the type specimens and cultivated plants. Species limits especially of U. mairei, U. fissa and U. himalayensis have been consistently confused in both herbaria and floristic treatments. As part of a taxonomic revision we produced a molecular phylogeny of the group based on ITS1-5.8S-ITS2, psbA–trnH, trnL–trnF and trnS–trnG markers. Our revision recovers a total of five species and two subspecies: U. fissa E.Pritz. ex Diels, U. grandidentata Miq. subsp. grandidentata, U. grandidentata Miq. subsp. lombok K.Becker & Weigend, U. himalayensis Kunth & C.D.Boché, U. mairei Lév. and U. parviflora Roxb. We provide type information, diagnostic characters, a key to species identification, species conservation assessments and a list of exsiccatae for each taxon.
... 4. Boraginaceae (Boraginoideae, tribe Boragineae): The stem node of Moritzia was set the same priors as described for Cryptantha as these fossils were also reported in the Ogalalla formation (Thomasson, 1979;Gabel et al., 1998). In this case the stem node of Moritzia was constrained following Weigend et al. (2010) (Chandler, 1961(Chandler, , 1962(Chandler, , 1964, which are the oldest fossils assigned to this family. 7. Heliotropiaceae: The stem node of Tournefortia L. sect. ...
... In our analyses, only Brazilian accessions of Moritzia are included, but the current geographic distribution of the sister genus Thaumatocaryon in South America indicates that this clade reached South America after a LDD event originating from Western Europe (Figures 1, 2). The abundance of fossils assignable to this clade in Miocene deposits of Kansas suggests that the MRCA ancestor of Moritziinae dispersed first to North America and from here to South America, going extinct in North America afterwards (Weigend et al., 2010). Without samples of Thaumatocaryon the exact time of this dispersal was, however, not possible to infer. ...
Long-distance dispersal seems to be the main biogeographic event responsible for intriguing distribution patterns in plant groups in which sister taxa are separated by thousands of kilometers of distance across oceans and continents. The biotic and abiotic mechanisms behind such dispersal events are poorly understood and many attempts have been made to explain how plants can manage to disperse and survive these long journeys. The biogeographic history of Boraginaceae, a subcosmopolitan plant family with many disjunct clades, is here addressed and analyzed in the context of the different dispersal modes exhibited by the species. The lack of a clear pattern between the main dispersal events in Boraginaceae and the phylogenetic distribution of the dispersal modes indicates that no single dispersal mechanism can be associated with the events of dispersal in the family. Moreover, adaptations to different dispersal agents and unassisted dispersal modes in some clades might have promoted the diversification of Boraginaceae in various habitats across several continents. Our study reveals that long-distance dispersal is a very complex process that needs to be analyzed in the context of climatic and environmental changes and the response of plants and their dispersal vectors to these variable conditions.
... Relazioni filogenetiche tra i generi della tribù Boragineae (albero semplificato da WEIGEND & al. 2010. In grassetto i nomi dei taxa presenti in Italia. ...
... This extremely variable fruit morphology of the Cynoglosseae has grossly misled genus-level taxonomy. Recent molecular studies show that some of the most striking carpological characters, traditionally considered as apomorphies defining genera, are actually highly homoplasious (Långström & Chase 2002;Thomas et al. 2008;Hasenstab-Lehman & Simpson 2012;Weigend et al. 2009Weigend et al. , 2010Weigend et al. , 2013Cecchi & Selvi 2009Coppi et al. 2015. Omphalodes is one of these polyphyletic genera (Weigend et al. 2013). ...
Molecular studies have recently led to major changes to the systematics of Boraginaceae and changed our perception of fruit morphology, previously used as the most important character in Boraginaceae classification. Navel-shaped nutlets as the key character for Omphalodes turned out to be highly homoplasious, thus the genus was revealed to be polyphyletic. After the recent reclassifications, the genus Omphalodes is still left paraphyletic, since the remaining taxa correspond to two different lineages: a clade of essentially Mediterranean perennials including the type species (O. verna) and a clade with the annual to subperennial North American taxa including the genus Mimophytum. We here advocate transferring all taxa of the North American clade into the genus Mimophytum, subdividing the genus Omphalodes into morphologically coherent and monophyletic entities. Mimophytum was initially removed from Omphalodes based on its glochidiate nutlets, but this character state does not define a monophyletic entity. In overall morphology, the redefined genus Mimophytum is highly homogeneous and easily differentiated from the other clades: it has a lax growth habit, and leaves are long-petiolate, abruptly contracted at the base into a more or less cordate base, and with a fine pubescence of erect trichomes. After the proposed rearrangements, only O. erecta remains as a problematic species whose systematic placement is still to be found out.
