The alien Solanum species in European Russia and the North Caucasus have never been revised. Over the past three decades, several exotic species have been found in various parts of the area, but many of them were identified incorrectly or reported using outdated taxonomy. We detailed the distribution data for these species, verified and corrected published reports, and provided updated diagnostic characters and a new identification key. Here, we report and discuss the records and residence status of nine species: S. elaeagnifolium, S. emulans, S. heterodoxum, S. nitidibaccatum, S. rostratum, S. scabrum, S. sisymbriifolium, S. triflorum, and S. villosum. One additional species, S. nigrum, appears to be an archeophyte widely distributed in both regions, except the Arctic zone. Solanum villosum, previously treated as several segregate taxa (S. alatum, S. luteum, S. transcaucasicum, S. zelenetzkii), was first collected in the early 20th century in the North Caucasus, whereas its collections from European Russia are mostly from the late 20th century. Solanum rostratum is naturalized in the plains of the North Caucasus and the Lower Volga Region. It has also been found several times in the forest and forest-steppe zones, with one new record reported here. Solanum triflorum is naturalized and rapidly spreading in the Lower Volga and the eastern North Caucasus; it is recorded here as a new casual alien in Kazakhstan. Solanum nitidibaccatum is reported for the first time from several administrative units and is considered naturalized at least in Central Russia. The putative presence of its close relative, S. sarrachoides from South America, which is a naturalized alien in some parts of Europe, is also discussed. At present, the remaining species are considered casual aliens. Solanum scabrum has escaped from cultivation, whereas the other species have arrived with contaminated grain imported from North America.
A checklist of the vascular flora of the Democratic Republic of the Congo (DRC) has been assembled, based on literature, online databases, and herbarium collections. It includes 10,260 species (38 lycophytes, 347 monilophytes, 16 gymnosperms, and 9,859 angiosperms), belonging to 246 families. The five largest families are Fabaceae (11.8% of specific richness), followed by Rubiaceae (6.7%), Asteraceae (6.6%), Poaceae (6.1%), and Orchidaceae (5.6%). Strict endemics of the DRC amount to 1,099 species, i.e., 10.7%. Families represented by > 120 species and comprising a high proportion of endemics include Commelinaceae (endemism rate: 24%), Euphorbiaceae (21%), Asteraceae (16%), and Fabaceae (16%). A total of 461 introduced species are recorded outside cultivation in the country, i.e., 4.5% of the total flora. Solanaceae, Brassicaceae, and Amaranthaceae stand out with their higher-than-average proportions of introduced species. Herbs represent the most frequent life form (54%), followed by (sub)shrubs (17%), trees (14.5%), climbers (10%), and epiphytes (4%). The IUCN conservation status was retrieved for 2,633 species (25.7%), of which two species are extinct and 448 (17%) are assigned to a threat category (CR [Critically Endangered]: 36; EN [Endangered]: 184; VU [Vulnerable]: 228). IUCN status was retrieved for 184 (17%) of the endemic species, of which 135 (73%) are considered extinct or threatened.
Limeum is the sole genus of Limeaceae, comprising approximately 20 species distributed across Africa, Southwest Asia, and India. However, genomic resources for the family are limited, and phylogenetic relationships within the genus remain poorly understood. In this study, we sequenced, assembled, and analyzed the complete chloroplast genomes of six Limeum species to investigate their genomic characteristics and phylogenetic relationships. The chloroplast genomes ranged from 156,354 bp to 160,377 bp in length and exhibited the typical quadripartite structure. Each genome contained 131 genes, consisting of 86 protein-coding genes, 37 tRNA genes, and eight rRNA genes. Comparative analyses identified 455 tandem repeats and 616 simple sequence repeats, revealing 16 highly variable regions that may serve as potential molecular markers for future studies. Phylogenetic analyses based on complete chloroplast genome sequences strongly supported the monophyly of Limeum and robustly resolved interspecific relationships within the genus. These results provide the first comprehensive plastome dataset for Limeum and a valuable genomic resource for species identification, genetic diversity assessment, phylogenetic reconstruction, and evolution studies of Limeum and the family Limeaceae.
Chenopodiaceae s.s. (Amaranthaceae s.l.) contains the largest number of C4 species among eudicots. Despite this, plastome evolution within this family has been investigated in only a few species. Here, we analyzed 119 plastomes from 115 species, including 78 newly sequenced plastomes, representing all subfamilies and most C4 lineages of Chenopodiaceae s.s. Plastome structural variants, rearrangements, and codon usage bias were compared across subfamilies and photosynthetic types. Multiple phylogenetic approaches were employed to reconstruct the evolutionary relationships within Chenopodiaceae s.s., and Bayesian divergence time estimation was performed. Various Mk models for discrete character evolution were tested to investigate the evolution of C4 photosynthesis, and stochastic character mapping simulations were used to reconstruct shifts in photosynthetic pathways through time. Several plastome structural variants and rearrangements were identified, but associations with photosynthetic types were observed only in the subfamily Suaedoideae. Codon usage bias analysis revealed significant bias exclusively in C4 species, suggesting enhanced translational efficiency and accuracy as an adaptation to environmental conditions. We inferred multiple independent origins of the C4 pathway, with the oldest lineages-Bienertia (Suaedoideae) and Caroxyleae (Salsoloideae)-dating to approximately 34 and 32 million years ago (Ma), respectively, during the Oligocene. A marked increase in the number of C4 lineages occurred between 20 and 15 Ma. Declining atmospheric CO2 concentrations, combined with genetic, ecological, and environmental factors, likely promoted the expansion of C4 photosynthesis until recently. Finally, we identified five new hypervariable regions that will be valuable for phylogenetic and DNA barcoding applications in Chenopodiaceae s.s.
Bassia scoparia is a widespread weedy species in the temperate regions of the world and is valued as a medicinal and ornamental plant. To date, the taxonomic concept of B. scoparia remains insufficiently studied due to a limited number of samples used in the previous phylogenetic analyses. To solve the taxonomy of the B. scoparia complex, we constructed a new phylogeny based on the nuclear ribosomal internal transcribed spacer (ITS), plastid intergenic spacer atpB-rbcL, and plastid region rpL16 intron sequences for numerous samples with diverse morphology. Our analysis revealed a close proximity and intermixed positions of the samples of the B. scoparia group with various morphology. Because of this polyphyly, we prefer to broadly delimit the species. An updated nomenclature of B. scoparia is provided including four new synonyms: Bassia angustifolia, B. littorea, Kochia albovillosa, and K. scoparia subsp. hirsutissima. In its new circumscription, B. scoparia encompasses populations with glabrous or variously hairy leaves and perianths. The original material of Kochia sieversiana, previously considered a species with hairy leaves and inflorescences, has the same diagnostic characters as in B. scoparia s.str. The correct name for more hairy-leaved plants is B. scoparia var. subvillosa. Plants with hairy perianths known as Kochia albovillosa and K. scoparia subsp. hirsutissima have a restricted distribution in Central Asia and South Siberia and have never been recorded as alien in other regions; they can be classified as a separate variety, B. scoparia var. hirsutissima. The ornamental variant of oblong or pyramidal shape may be called B. scoparia var. trichophila. Bassia scoparia is often confused with a similarly looking relative, B. indica, especially in North Africa, a region where secondary ranges of both species overlap. Phylogenetically, these species are sister groups; they share some morphological characters but have different primary distribution ranges. We traced a recent expansion of B. indica in the Mediterranean with the first record reported from the European continent (Spain) and uncovered various introduction pathways of the species in this region.
Desmochaeta (now Cyathula) achyranthoides was described from South America and reported to be a widespread tropical plant in both Africa and the Americas. A revision of herbarium material revealed that inter alia leaf shape differs between the populations of the Eastern and Western Hemispheres. Therefore, we maintain the name C. achyranthoides s.str. for the American populations and re-instate the name C. geminata for most of the African plants. Both species are found in tropical evergreen forests, mainly at low altitudes. Furthermore, two mountain species, C. brevispicata from Madagascar and C. aethiopica from east tropical Africa, which were previously identified as C. achyranthoides, are described as new to science. Compared to both C. achyranthoides and C. geminata, these new species have short inflorescences and longer, recurved or uncinate perianths in the fertile flowers and morphologically resemble C. fernando-poensis; the latter is only known from the mountains of Equatorial Guinea (Bioko Island), south-west and (newly recorded here) North-West Regions of Cameroon. The species under study are compared with one another and with the related, pantropically distributed species C. prostrata; their synonymy is verified and typifications are established. The fine-level partial florescence (cyme) structure of each species is also studied, with further taxonomic implications. Cyathula geminata seems to be restricted to west and central tropical Africa, with its range replaced eastwards by C. aethiopica and in Madagascar by C. brevispicata.
The application of omics data serves as a powerful tool for investigating the roles of incomplete lineage sorting (ILS) and hybridization in shaping genomic diversity, offering deeper insights into complex evolutionary processes. In this study, we utilized deep genome sequencing data from 76 individuals of Lappula and its closely allied genera, collected from China and Central Asia. By employing the HybPiper and Easy353 pipelines, we recovered 262-279 single-copy nuclear genes (SCNs) and 352-353 Angiosperms353 genes, respectively. We analyzed multiple datasets, including complete chloroplast genomes and a filtered set of 475 SCNs, to conduct phylogenetic analyses using both concatenated and coalescent-based methods. Furthermore, we employed Quartet Sampling (QS), coalescent simulations, MSCquartets, HyDe, and reticulate network analyses to investigate the sources of phylogenetic discordance. Our results confirm that Lappula is polyphyletic, with L. mogoltavica clustering with Pseudolappula sinaica and forming a sister relationship with other taxa included in this study. Additionally, three Lepechiniella taxa nested within distinct clades of Lappula. Significant gene tree discordance was observed at several nodes within Lappula. Coalescent simulations and hybrid detection analyses suggest that both ILS and hybridization contribute to these discrepancies. Flow cytometry (FCM) analyses confirmed the presence of both diploid and tetraploid taxa within Lappula. Phylogenetic network analyses further revealed that Clades IV and VII likely originated through hybridization, with the tetraploids in Clade IV arising from two independent hybridization events. Additionally, the "ghost lineage" identified as sister to Lappula redowskii serves as one of the donors in allopolyploidization. In conclusion, our study provides new insights into the deep phylogenetic relationships of Asian Lappula and its closely allied genera, contributing to a more comprehensive understanding of the evolution and diversification of Lappula.
Achyranthes in its traditional sense (excluding Achyropsis that phylogenetically falls into Achyranthes) has been considered to contain a restricted (three to four) number of species in Africa and one or two species in the Arabian Peninsula. The morphology of the type species of the genus, A. aspera, has been treated as highly polymorphic, with several varieties recognised by various authors. Not surprisingly, a recent extended phylogeny revealed a non-monophyly of A. aspera. We present a deeper insight into morphological characters of the A. aspera aggregate together with taxonomic, nomenclatural, ecological, and chorological data based on field investigations and herbarium studies. Instead of one polymorphic species, we accept A. aspera s.str., A. abyssinica, A. acuminata, A. annua, A. mauritiana, A. porphyrostachya, A. sicula, and A. seychellensissp. nov., all being native to different parts of Africa. In most herbaria, the vast majority of African specimens labelled as A. aspera belong to other species, which are being reinstated here. In addition, two well-recognized species, A. fasciculata and A. talbotii from Tropical East and West Africa, respectively, are also discussed. Moreover, we found that the type of A. aspera var. pubescens as listed in the African and Arabian floras and checklists belongs in fact to an American species A. fruticosa, which is absent in the Old World. In place of the misapplied A. aspera var. pubescens, we accept A. porphyrostachya, a species described from Myanmar, as a correct name for the populations growing in Africa and Arabia. According to our results, at least 10 native species of Achyranthes occur in Africa (or 16 species if Achyropsis is merged with Achyranthes), which is a major diversity center of the genus. Four species are recorded from the Arabian Peninsula (A. abyssinica, A. annua, A. aspera s.str., A. porphyrostachya), and two of them (A. abyssinica and A. annua) reach their easternmost range limit in this region. As a result, the distribution as well as ecological conditions of each species is now clarified or circumscribed for the first time.
Phlomoides bomiensis, a new species in Bomi County, Xizang, China, was described and illustrated. In addition, Phlomoides longidentata, previously only known from Nepal and Bhutan, is newly recorded from Dingri County, Xizang, China. The phylogenetic placement of both species within the genus was analysed using nine plastid DNA markers (atpB-rbcL, psbA-trnH, rpl16, rpl32-trnL, rps16, trnK, trnL-trnF, trnS-trnG, trnT-trnL). Both species have brown-black trichomes inside the upper corolla lip and nested within the same subclade of Clade II. A diagnostic key to the Phlomoides species belonging to this subclade is provided.