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We present the first taxonomic revision of Selaginella subg. Ericetorum on the basis of an investigation of all known species, using optical microscopy and scanning electron microscopy. Selaginella subg. Ericetorum is distinguished from other Selaginella subgenera by the presence of the combination of monomorphic and decussately arranged leaves. Previous studies distinguished three species, namely, S. gracillima and S. uliginosa from Australia, and S. pygmaea from Australia and South Africa; our investigation revealed five species. Because of morphological differences and geographical separation, we proposed segregation of S. aboriginalis (of Australia) from S. pygmaea (of South Africa). The fifth species, S. royenii from Papua New Guinea, was described in 1956 but remained unnoticed. Although there are only five species, these are frequently misidentified in herbarium collections because of a lack of a monographic treatment. Therefore, in this study, we present two identification keys using different macro- and microstructural characters facilitating the identification of all the species. In addition, we present species descriptions and photographs showing structural details. For the first time, it is shown that the sculpture of Selaginella megaspores has sufficient diversity to distinguish closely related species while maintaining a high level of uniformity within a species.
Polyploidy, hybridization, and agamospory have been considered important mechanisms in fern speciation. By integrating the methods of cytology, molecular phylogeny, and morphology, we examined the origins of polyploid species in the Pteris cretica group, which comprises five agamosporous taxa and six sexual species. Phylogenetic analysis was conducted using both cpDNA (rbcL and trnV-trnM) and a low-copy nuclear gene (gapCp). The combined results of cytology and the phylogenetic trees suggested that the sexual diploid P. kidoi had played a central role in the diversification of polyploid species in the P. cretica group. Some triploid clones of agamosporous P. cretica var. cretica originated through hybridization between the diploid cytotype of P. cretica var. cretica and the sexual diploid P. kidoi. The sexual polyploid species, P. henryi, P. multifida, P. ryukyuensis, and P. yamatensis, and the hexaploid race of P. deltodon have arisen through allopolyploidization between their respective ancestral parent species and P. kidoi. Additionally, the agamosporous triploid P. nipponica and P.cretica var. albolineata might have originated through hybridization between P. cretica var. cretica and an unknown ancestral diploid parent of P. ryukyuensis. Agamosporous P. cretica var. cretica harbored considerable genetic variation within both the diploid and triploid cytotypes. Because we could not find clear genetic differences between the diploid and triploid P. cretica, both cytotypes might be autopolyploids, or alternatively have originated through hybridization among the same members of ancestral sexual species.
Adiantum lindsaeoides (Pteridaceae) is a new fern species from the Atlantic Rain Forest, on the coast of Brazil. It can be recognized by the nodose rhizomes, stipes approximate, dense indument of short hairs on both sides of the stipes and rachises, plus scattered, slightly tortuous, filiform scales, and by its median segments conspicuously falcate, and dimidiate that resemble the segments of a Lindsaea. The most similar species to A. lindsaeoides is A. mynsseniae that differs by having rhizomes not nodose, stipes apart, stipes and rachises glabrous abaxially, and segment apices long-acuminate to acute (vs. round in A. lindsaeoides). So far, A. lindsaeoides is only known from Bahia and Espírito Santo States. The new species is described and illustrated, and, a key to the most similar morphological species is given.
The Schismatoglottid Alliance (tribes Schismatoglottideae, Cryptocoryneae and Philonotieae: Araceae) has been shown to be monophyletic in a previous study based on two cpDNA regions (matK and trnL-F). Here, ten additional taxa of Schismatoglottideae were sequenced, including two extra-Bornean Piptospatha species, and the type species of Hottarum, unavailable at the time of the previous analyses. Phylogenetic analyses were performed by using parsimony, likelihood (RAxML and likelihood ratchet PAUP*), and Bayesian inference to determine the placement of these additional taxa within Schismatoglottideae. The resulting tree topology supports, 1) a monophyletic West Sarawak clade comprising three genera: Aridarum, Bakoa, and Piptospatha; and 2) a supra-Lupar Line (the boundary between two of the known Bornean biochores) clade comprising the ‘Schismatoglottis’ josefii complex. Nineteen morphological, ecological, and geographical characters were coded and mapped using parsimony (unordered model) onto the Bayesian tree to investigate incidents of homoplasy or apomorphic status of morphologies presumed significant in the evolution of rheophytism. The morphologies proposed here as the primary mechanisms adaptive for rheophytism in Bornean Schismatoglottideae are: root/shoot disarticulation; a free ligular sheath; and marcescent senescence of the ligule. Two morphologies formerly used as generic definers, and previously treated as homologous: presence of a micropylar appendage, and thecae horns, are shown to be homoplastic.
We demonstrate, through separate and combined phylogenetic analyses of morphological characters, nucleotide sequences, and coded gaps of two DNA regions (plastid rpl32-trnL and nrDNA-ITS) using parsimony and Bayesian inference, that Lophiarella (Orchidaceae), as originally circumscribed, is polyphyletic. We re-circumscribe the genus to exclude one of the two species included in the original description and include two others, which requires two new combinations: Lophiarella splendida and Lophiarella flavovirens. As newly circumscribed, Lophiarella is strongly supported as a monophyletic assemblage within the Trichocentrum-clade, where it is sister to the (Cohniella, (Trichocentrum, Lophiaris)) clade. Lophiarella is easily diagnosable within the Trichocentrum-clade, among other characters, by the conduplicate yet rigidly fleshy-coriaceous leaves and the rigidly erect inflorescences with peduncles coated with a conspicuous, waxy film. In the context of a phylogenetic analysis, we discuss the evolution of several taxonomically, ecologically and biogeographically relevant characters, such as rostellum shape, vegetative architecture, labellum, and column structure. We also present a key to the genera of the Trichocentrum-clade and a taxonomic revision of the species of Lophiarella with keys, descriptions, nomenclature, iconography, discussion of affinities, ecological and biogeographical information, a distribution map, and full exsiccata citations.
Cremastra malipoensis G. W. Hu, a new rare species of Orchidaceae, is described and illustrated, and a taxonomic key to the species of the genus is provided. It occurs in southeastern Yunnan, China, near the border of Vietnam. It is most similar to C. appendiculata, but its inflorescences have fewer flowers (only 4–7), its flowers are positioned horizontally and become slightly pendulous, the lateral lobes of the lip are geniculate and twisted 90° at the base, and the mid-lobe of the lip is somewhat reflexed, rhombic to broadly ovoid, and 9–10 × 8–9 mm. The new species is known only from a single population and categorized under the ICUN criteria as critically endangered. Formal recognition is required for protecting this species, which grows sympatrically with more common species that are harvested as medicinal herbs.
Irrespective of the fact that Borneo is one of the richest areas for mycoheterotrophic plants, only two species of the genus Lecanorchis Blume, a mycoheterotrophic genus of Orchidaceae, have been reported from Borneo. On three occasions during recent botanical surveys in Betung Kerihun National Park, we found an undescribed species of Lecanorchis. Here, we provide a detailed morphological account of this new species, Lecanorchis betung-kerihunensis. We also provide a key to the species of Lecanorchis in Borneo.
Tillandsia bonita is described as a new species and is illustrated. It is compared with Tillandsia didisticha, which we assume is the morphologically most similar species. The new species is the ninth taxon of this genus recorded for the flora of the state of Mato Grosso do Sul, central-western Brazil. Illustrations and a brief discussion on the leaf anatomy of the species are provided. Two additional species that have been overlooked by previous authors are included in the list of Bromeliaceae of Mato Grosso do Sul and an updated key for Tillandsia from that state is presented.
Divergence between evolutionary lineages is not always marked by the development of obvious species-specific characters, whether morphological, physiological, or ecological. Consequentially, extant biodiversity can easily be overlooked. These cryptic species are often not recognized until genetic data are in hand, as is the case for the novel taxon we describe here. Carex viridistellata in Carex section Ceratocystis is an endemic species restricted to calcareous wetlands of Michigan, Ohio, and Indiana, where it has previously been collected as Carex cryptolepis. Crosses between Carex viridistellata and Carex cryptolepis produce sterile F1 hybrids, and the two species are differentiated by a number of subtle morphological characters, as well as aspects of their respective ecologies. Phylogenetic analyses of nrDNA strongly indicate monophyly of Carex viridistellata and its sister species relationship with the North Carolina narrow endemic Carex lutea. Both species constitute a distinct lineage within a generally poorly resolved section Ceratocystis. This work highlights the broad importance of cryptic taxa, with implications for fields ranging from population genetics to conservation and restoration.
Chusquea Kunth is the most diverse genus of bamboos with about 160 species accepted and an estimate of more than 50 to be described. This is an exclusively American genus with about 40 species recognized in Brazil, these occurring in montane grasslands or more commonly in montane forests. In Bahia State six species are known and a new species (Chusquea clemirae) is herein described, illustrated, and compared with sympatric or morphologically similar species. In addition, it is classified as a probable member of C. subg. Chusquea mainly based on internode length, shape and direction of the central bud, type of branching, and length and width of the foliage leaf blades. Its culms are slightly lignified with a spongy medulla and are solid to hollow; a vertically oriented central bud and other vegetative features clearly distinguish it from other Chusquea. This new species was found exclusively in a montane Atlantic Rainforest area in southern Bahia (the Serra Bonita Private Natural Heritage Reserve), and it has a restricted occurrence within the reserve.
Arthrostylidiinae are one of the three subtribes of Neotropical woody bamboos and comprise 13 genera and over 160 described species. Recent phylogenetic analyses have demonstrated both the non-monophyly of Aulonemia Goudot and the sister relationship of Aulonemia ulei (Hack.) McClure to the monotypic Glaziophyton Franch., with which it forms a clade sister to the reminder of the subtribe. Based on these data and on morphological and anatomical characters, we present herein the new monotypic genus Cambajuva, as well as a new combination, to accommodate A. ulei. Morphological and anatomical descriptions are also provided, as well as a comparison with putatively related genera of the subtribe.
We examined the phylogenetic relationships and patterns of character evolution among the California poppies and their relatives (Papaveraceae, Eschscholtzieae) using morphological comparisons and parsimony and likelihood-based phylogenetic analyses of two chloroplast and two nuclear DNA regions, including 14-3-3-like, a phylogenetically informative nuclear DNA region developed for this research. Each of the three genera in the tribe (Dendromecon, Eschscholzia, and Hunnemannia) is strongly supported as monophyletic, but not all species of Eschscholzia display monophyly based on molecular data. Examination of biogeographic patterns indicates that dispersal to island habitats occurred more than twice in the tribe. Phylogenetic reconstruction of the evolution of chromosome number suggests that there were several polyploidy events within the tribe and likely one or more aneuploid events within Eschscholzia. Based on the results presented here, the subspecies of both Eschscholzia californica and Eschscholzia lemmonii will be promoted to specific status. Additionally, the study provides evidence supporting recognition of two new species of desert Eschscholzia.
A new species of Lespedeza (Fabaceae) from Jiangxi Province, China is here described as Lespedeza jiangxiensis. The new species is morphologically similar to L. inschanica, but differs on the obovate leaflets with densely adpressed-pubescence adaxially and abaxially, shorter floral peduncles (0.5–1 mm long), shorter calyces (2.9–3.8 mm long), and white to yellowish-white keel-petal apices. In contrast, L. inschanica is characterized by oblong leaflets that are glabrous adaxially, longer floral peduncles (3–10 mm long), longer calyces (5–6 mm long), and light purple keel-petal apices. A phylogenetic analysis based on DNA sequences of plastid rpL32-trnL and trnL-F intergenic spacers identified L. pilosa as the closest relative of L. jiangxiensis. Interestingly, however, L. pilosa is remarkably different morphologically from L. jiangxiensis. Pollen grains of L. jiangxiensis are similar to those of L. chinensis, L. inschanica, and L. pilosa in size, shape, and germination organ type, but clearly different in exine sculpture.
A new species of “jurema,” Mimosa urandiensis (Leguminosae, Mimosoideae), from the municipality of Urandi, Bahia, is described and illustrated. This species is distinguished primarily by the presence of capitate glandular trichomes on vegetative and reproductive structures, associated with the glandular-ciliolate stipules and bracts. The morphological distinctiveness of this new species and its relationships with allied species are discussed. Scanning electron micrographs of trichomes and pollen grains, and a key to this new taxon and morphologically similar species are provided.
Pickeringia, a monotypic genus of the Leguminosae endemic to the xerophytic sclerophyllous chaparral vegetation of the California Floristic Province, has been considered a “relict” of legume diversity in the North American flora and an example of the classic “paleoendemics” in the flora of California. Evidence is presented for the phylogenetic relationships of this genus, inferred from nucleotide sequence variation in the internal transcribed spacers of nuclear ribosomal DNA and the plastid matK gene. Phylogenies derived from maximum parsimony and Bayesian analyses both strongly support a close relationship of Pickeringia to the temperate to subtropical, deciduous genera Cladrastis and Styphnolobium of tribe Sophoreae consistent with morphological and cytogenetic evidence rather than to members of tribe Thermopsideae where the genus has been treated taxonomically. Cladrastis is resolved as paraphyletic while Styphnolobium is strongly supported as a monophyletic group. These results, plus an estimated age of -31 million years for the genus, further substantiate the hypothesis that Pickeringia is geographically isolated in the flora of western North America, an old and phylogenetically distinct lineage of an early diverging group of papilionoid legumes that were much more widely distributed throughout temperate North America and Asia during the Tertiary but whose modern relatives are restricted to southern North America, Central America, and eastern Asia. These findings have implications not only for the evolutionary history of Pickeringia but also for the age and development of chaparral vegetation in the Californian flora.
A new species, Monnina goiana, is described based on a re-interpretation of the name M. martiana, and a key for the Brazilian species of Monnina is presented. The identity of M. martiana is discussed.
Disjunct distributions in poorly defined taxa can serve as indicators that the members of the isolated ranges are polyphyletic, as has been previously demonstrated in many plant families. Such taxa should be prioritized for inclusion in phylogenetic analyses so that at least one member from each isolated range should be sampled together with other closely related taxa. We describe a new monotypic genus (Wilczekra) for Euonymus congolensis, which is the only species of Euonymus reported in mainland Africa other than one other species (E. latifolius) whose range extends from Europe into northern Africa. We infer that Wilczekra congolensis is the sole African member of an early-derived lineage of Celastraceae that is most diverse in the Austral-Pacific but also includes Crossopetalum from the West Indies and tropical America.
Oxalis wulingensis, an unusual new species of Oxalidaceae from Hunan and Hubei provinces of central China, is described and illustrated. Placement of this species within Oxalis was assessed based on DNA analyses of the nuclear internal transcribed spacer (ITS) and morphological comparisons with related species. This new species can be easily distinguished by its long, obtriangular leaflets, the two lateral ones arranged at a 180° angle and densely pubescent with brown, villous hairs on both surfaces, pink flowers with lilac veins, and apex of petals obtuse or irregularly denticulate with 3–5 teeth. The chromosome number is 2n = 22. With the addition of O. wulingensis, which is the only species endemic to China, a total of seven native Oxalis occur in the country. A taxonomic key to all Chinese species of Oxalis is also provided.
Kuhlmanniodendron macrocarpum Groppo, Favaretto & Fiaschi (Lindackerieae, Achariaceae), a second species of the genus, is here described and illustrated. This species is known from a small number of collections from the state of Bahia, eastern Brazil, and considered endangered given its restricted geographical distribution and the rapid deforestation of the Brazilian Atlantic Rain Forest. Kuhlmanniodendron macrocarpum can be readily distinguished from K. apterocarpum, the other species of the genus, by its larger fruits (5–8 × 4–6 cm vs. 3–4 × 2.5–3 cm), the leaf venation pattern, and the disposition and number of stamens in male flowers. In addition to morphological characters used to characterize the new species, phylogenetic analyses were carried out using an rbcL sequence from the new species together with other Malpighiales sequences deposited in GenBank. These analyses involved maximum parsimony and Bayesian inference. The results confirmed the close relationship of the new species with K. apterocarpum in a strongly supported clade. Kuhlmanniodendron appeared close to Camptostylus, Lindackeria, Xylotheca, and Caloncoba in a strongly supported clade (posterior probability = 0.99) that corresponds with the tribe Lindackerieae of Achariaceae (Malpighiales). The arrangement of the stamens in the male flowers of the new species were investigated using transversal anatomical sections, revealing a combination of stamens with basally connate filaments externally arranged in a ring and internally arranged in sets of three (or two), the first report of this pattern of stamen arrangement in Achariaceae. Pollen of K. macrocarpum are 3-colporate and microreticulate, and typical of many eudicots. A picrosodic paper test was performed in herbarium dry leaves, and cyanogenic glycosides, a class of compounds usually found in Achariaceae, were detected. A detailed description and illustrations of this new species are provided, as well as comments on geographic distribution, phenology, generic relationships, and conservation status.
A new species of Serjania (Sapindaceae, Paullinieae), S. lucianoi from Northern Bahia, Brazil, belonging to Serjania sect. Eurycoccus, is described, illustrated, and compared to its putative closest relative. Serjania lucianoi seems to be related to S. glutinosa, and can be distinguished from it by: the 5-foliolate leaves, linear stipules, indument with simple and glandular obconical-headed trichomes, fruits with ovate outline, and seeds in basal portion of the locule. In addition, micromorphological characters of leaf and flower epidermis are given, and pollen grains are described. The new species is known only from Bahian caatinga, being an element of the dunes of the São Francisco River. We present a key to the Brazilian species of Serjania restricted to this biome.
Physarieae, a tribe of mustards with multiaperturate pollen, was investigated phylogenetically for the first time using comprehensive sampling and plastid molecular data. Phylogenetic analyses were employed to test the monophyly of the tribe, to investigate relationships of its seven genera, and to study the evolution of selected morphological traits. Character evolution was inferred using parsimony approaches for nine continuous and three discrete traits related to trichomes, fruits, seeds, and pollen. Two monophyletic clades were recovered within Physarieae: the DDNLS clade, which included five genera (Dithyrea, Dimorphocarpa, Nerisyrenia, Lyrocarpa, and Synthlipsis), and the PP clade, which comprised two genera (Paysonia and Physaria). Under this new phylogenetic context, multiaperturate pollen appeared to be a potential synapomorphy of the tribe. In addition, five continuous fruit and seed traits were identified as new potential synapomorphies for Physarieae. Our analyses also indicated that members of the DDNLS clade present more highly branched dendritic trichomes, narrower replums, longer fruits, and wider seeds, when compared to representatives of the PP clade. Moreover, ancestral state reconstructions of continuous traits in Physarieae showed that seeds became wider and fruits, replums, styles, and seed primary grooves became longer over time.
Boechera (Brassicaceae) is a diverse genus of ±70 sexual diploid species and numerous apomictic hybrids concentrated in western North America. It is the largest genus in tribe Boechereae, which also includes seven other small genera. Boechera is closely related to Arabidopsis and is becoming a model genus for diverse studies, focusing particularly on apomixis and hybrid speciation. As part of an ongoing effort to clarify the taxonomy of the group, we present phylogenetic analyses of Boechereae, including all genera of the tribe and most of the sexual diploid species of Boechera. Ten loci are used, including two plastid loci, nrDNA ITS, and seven low-copy nuclear loci. These analyses indicate that Boechera, as currently circumscribed, is polyphyletic, comprising three main clades. Eastern North American species previously assigned to Boechera form a distinct clade with the Asian taxon Borodinia macrophylla and are herein transferred to the genus Borodinia, resulting in seven new combinations (Borodinia burkii, Borodinia canadensis, Borodinia dentata, Borodinia laevigata, Borodinia missouriensis, Borodinia perstellata, and Borodinia serotina). Boechera repanda, a morphologically aberrant species endemic to the Sierra Nevada, is recognized in the new genus Yosemitea (as Yosemitea repanda). A primarily western North American clade comprising all remaining species is strongly supported and retains the name Boechera. Although resolution within Boechera sensu stricto is limited, a number of well-supported clades are identified.
The genus Harrisia (Cactaceae) comprises 20 species native to South America and the Caribbean. To investigate relationships within the genus, plastid and nuclear markers were sequenced from Harrisia and related genera. Phylogenetic analyses recover a monophyletic Harrisia that can be divided into two subgenera, four sections, and two series. Harrisia likely originated in the Andes and later dispersed into Brazil before further speciation in the Gran Chaco or the Caribbean. A key is given for the taxonomic classifications proposed for Harrisia. New combinations are H. subg. Eriocereus and H. sect. Roseocereus. New taxonomic names are H. sect. Adscendens and H.ser. Earlei.
This paper presents descriptions and illustrations for two new species of Gentianaceae, Tripterospermum maculatum from Sichuan Province (China) and T. tanatorajanense from Sulawesi (Indonesia). Based on literature search, Tripterospermum maculatum is morphologically close to T. pingbianense and T. lanceolatum, and T. tanatorajanense resembles T. luzonense and T. alutaceifolium. To test if the new species differ from their morphologically most similar species, we measured various traits on herbarium specimens and performed a principal component analysis (PCA). This analysis showed that the new species differ from similar species in gross morphology for several diagnostic traits. Tripterospermum maculatum differs from T. pingbianense and T. lanceolatum by having calyx lobes longer than calyx tubes, a whitish-yellow corolla maculated with purple dots, and a gynophore shorter than the ovary. Tripterospermum tanatorajanense is distinct from T. luzonense by having a narrowly winged calyx. It differs from T. alutaceifolium and T. luzonense by having a shorter ovary and a slightly longer gynophore. A key including all species of Tripterospermum is provided.
Gardenia is an Old World genus of about 200 species. A bibliographic survey on Sulawesi Gardenia, a region extremely rich in plant diversity, but poorly explored botanically, enumerated five species, namely G. affinis, G. forsteniana, G. mutabilis, G. pelenkahuana, and G. vernicosa. Of these, G. mutabilis remains in Gardenia based on current generic delimitation using various morphological characters of the vegetative and floral parts. Three taxa, G. affinis, G. forsteniana, and G. pelenkahuana, have been reassigned to genera such as Porterandia or Rothmannia. Meanwhile, G. vernicosa was found to have been erroneously listed for Sulawesi. As herbarium materials labeled as G. mutabilis were examined and sorted, they were found to be heterogeneous and included two novelties, G. kabaenensis and G. longistipula, described and illustrated here. These two new taxa are distinguished from G. mutabilis based on calyx form and tube length, and stipule length.
A new concept of the genus Heterocoma is presented based on morphological and anatomical evidence. The genus is characterized by the combination of foliose subinvolucral bracts, fimbrillate or paleaceous receptacles, glabrous cypselae, and a caducous biseriate pappus (at least the inner series). A putative synapomorphy of the genus is the presence of phytomelanin in the cypselae walls. The monotypic genera Bishopalea, Sipolisia, and Xerxes are synonymized under Heterocoma with the following new combinations: Heterocoma ekmaniana, Heterocoma erecta, and Heterocoma lanuginosa. Two new species, Heterocoma gracilis and Heterocoma robinsoniana, are described and illustrated. The genus is endemic to the campos rupestres of the Brazilian Central Plateau. A key to the six species recognized in Heterocoma is provided.
Analysis of DNA sequence data was used to clarify the circumscription and relationships of the small Mexican genus Asanthus. Results from non-coding regions, including the nuclear ITS and the chloroplast psbA-trnH spacer showed that Asanthus is clearly monophyletic and distinct from Steviopsis and Brickellia, two genera with which it is sometimes lumped, as well as from the closely related Brickelliastrum and Carminatia. The DNA sequence data also agree with morphological observations in showing that Asanthus is composed of three species, A. thyrsiflorus from the Sierra Madre Oriental and A. squamulosus and A. solidaginifolius from the Sierra Madre Occidental. Samples from the southern Sierra Madre Occidental that have been identified as A. thyrsiflorus are suggested by the presence of an additive pattern of polymorphisms in the nuclear ITS region to be of interspecific hybrid origin between A. thyrsiflorus and A. solidaginifolius, which is consistent with reports of their morphological intergradation in this area. The results suggest that geographic separation and divergence within Asanthus is now being followed by secondary contact, and point to a dynamic pattern of movement and change in the genus.
Gamochaeta beckii, a new species found in La Paz, Bolivia is described and illustrated. Gamochaeta beckii is similar to G. purpurea but it is easily distinguished by stems branched from the base to the inflorescence, oblanceolate leaves, the small capitula arranged in glomerules, forming leafy short and continuous spikes, and short acuminate inner phyllary apices. A key to the species of Gamochaeta occurring in Bolivia is presented.
Classification of Angelica and its allies is complex and controversial, with previous phylogenetic studies restricted to examination of small numbers of taxa using only nrDNA ITS sequences. In this study, phylogenetic analyses of nrDNA ITS and ETS sequences, cpDNA sequences (rps16 intron, rps16-trnK, rpl32-trnL, and tmL-trnT), and morphological data, supplemented by observations of fruit anatomy and micromorphology, were used to ascertain evolutionary relationships and confirm generic boundaries within Angelica s.l. (including Angelica, Archangelica, Coelopleurum, Czernaevia, and Ostericum), with emphasis on its East Asian members. Most species of Angelica s.l. fall into two major, disparate clades, the Ostericum clade and the Angelica group, with the latter comprising five major lineages that are distinguished both molecularly and morphologically: Angelica s.S. (including Czernaevia), Archangelica, Coelopleurum, Glehnia, and a newly identified Littoral Angelica clade. A North American Angelica clade was also distinguished in the ITS trees. Taxonomic realignments will be required, as many species of Angelica fall outside of Angelica s.S. and four species of Angelica occur outside of the Angelica group (A. hirsutiflora, A. oncosepala, A. paeoniifolia, and A. sinensis). The Angelica s.S. clade contains predominantly East Asian species and comprises five major lineages, two of which represent plants exclusively from the eastern Himalayas; each of these five lineages can also be defined morphologically. The results obtained are significantly different from traditional treatments of Angelica s.l. and provide new insights into the phylogeny and classification of the group.
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