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Adiantum meishanianum is an endemic species distributed only in Meishan village, Kaohsiung, Taiwan. Because its sporangia contain only abortive spores, A. meishanianum has been regarded as having a hybrid origin, presumably with A. caudatum, A. malesianum, and A. philippense as the putative parents. The aim of this study is to confirm the hybrid origin and determine the parental lineages of A. meishanianum by examining cytology, reproductive modes, and using molecular phylogenetics. We found that the sequences of two chloroplast regions, rps16-matK intergenic spacer and the matK gene, are identical between A. meishanianum and A. malesianum, suggesting A. malesianum is the maternal parent. The nuclear phylogeny reconstructed based on the low-copy marker, CRY2 first intron, reveals that A. meishanianum has three types of sequences: one type groups with sequences of sexual diploid individuals of A. philippense and the other two group with sequences of the sexual tetraploid A. malesianum, indicating that A. philippense is the paternal species. Our data further imply that the extant A. meishanianum probably originated from a single hybridization event, and its rarity is likely due to the limited distribution of the paternal parent.
The Polypodium vulgare complex (Polypodiaceae) comprises a well-studied group of fern taxa whose members are cryptically differentiated morphologically and have generated a confusing and highly reticulate species cluster. Once considered a single species spanning much of northern Eurasia and North America, P. vulgare has been segregated into 17 diploid and polyploid taxa as a result of cytotaxonomic work, hybridization experiments, and isozyme studies conducted during the 20th century. Despite considerable effort, however, the evolutionary relationships among the diploid members of the P. vulgare complex remain poorly resolved. Here we infer a diploids-only phylogeny of the P. vulgare complex and related species to test previous hypotheses concerning relationships within Polypodium sensu stricto. Using sequence data from four plastid loci (atpA, rbcL, matK, and trnG-trnR), we recovered a monophyletic P. vulgare complex comprising four well-supported clades. The P. vulgare complex is resolved as sister to the Neotropical P. plesiosorum group and these, in turn, are sister to the Asian endemic Pleurosoriopsis makinoi. Using divergence time analyses incorporating previously derived age constraints and fossil data, we estimate an early Miocene origin for the P. vulgare complex and a late Miocene-Pliocene origin for the four major diploid lineages of the complex, with the majority of extant diploid species diversifying from the late Miocene through the Pleistocene. Finally, we use our node age estimates to reassess previous hypotheses, and to propose new hypotheses, about the historical events that shaped the diversity and current geographic distribution of the diploid species of the P. vulgare complex.
Ricardo S. Couto, Vitor Tenorio, Fernanda da C. Alzer, Rosana C. Lopes, Ricardo C. Vieira, Cláudia B. F. Mendonça, Vania Gonçalves-Esteves, João Marcelo Alvarenga Braga
The Dioscoreaceae are widely distributed in the tropics, but despite the diversity and utility of the taxa of this family as food (e.g. yam) they are often poorly delimited morphologically and have complicated taxonomic issues. The Dioscorea campestris assemblage comprises 11 names in the literature, with considerable morphological variation among herbarium specimens. Macro- and micro-morphological examinations of the tubers, stems, and pollen indicated the recognition only of D. campestris and the proposal of a new combination— Dioscorea pedalis; the other nine names are reduced to synonyms of D. campestris. We report here the pollinic, anatomical, and morphological characters of D. campestris and D. pedalis, and describe the ecology, phenology, geographical distributions, etymology, conservation status, and taxonomies of both species.
A new species of Anathallis, A. roseopapillosa, is described from the central Brazilian Amazon. It is morphologically related to A. gracilenta, but differs by having shorter racemes arising abaxially from the leaves, with flowers opening one at a time, petals with a papillose apex, and an oblong-elliptic to ovate-elliptic lip, which is often obscurely tri-lobed. We also provide evidence of the phylogenetic position of the new species in the subtribe Pleurothallidinae by means of a cladistic analysis of nuclear DNA sequence data (ITS).
A new orchid species, Encyclia sabanensis, is described and illustrated. The new taxon is compared with a large group of species of Encyclia from the Caribbean. The distribution, ecology, and conservation status of E. sabanensis are briefly discussed. A general field key for identifying 24 Encyclia species that has been reported for Cuba is offered. The key is coupled with the diagrams of the labellum of 23 species.
Six different morphotypes for the Scilla species in western Europe have been distinguished based on leaf width, length of the basal bract, number of flowers per inflorescence, and length of bulbs: (1) S. peruviana; (2) S. hyacinthoides; (3) S. lilio-hyacinthus; (4) S. rambureiS. beirana; (5) S. vernaS. pauiS. odorataS. merinoiS. monophyllos; and (6) S. obtusifoliaS. numidicaS. autumnalis. Two biogeographic trends have been circumscribed in terms of monthly precipitation averages (the most discriminating parameter). One trend, characterized by lower annual regimes of precipitation and distributions around the Mediterranean basin, is represented by S. peruviana, S. hyacinthoides, S. obtusifolia, and S. numidica. The second trend, associated with higher average precipitation along the extreme western coast of Europe, is represented by the S. verna group and S. lilio-hyacinthus. Scilla autumnalis exhibits an intermediate environmental behaviour. Future climate change scenarios point towards an increase in the potential habitat area of species distributed along the Mediterranean basin (S. peruviana, S. hyacinthoides, S. obtusifolia, S. numidica, and S. autumnalis) and a decrease in the potential habitat area for the western-most likely morphotypes (S. ramburei, S. verna, S. monophyllos, and S. lilio-hyacinthus).
Over one third of African palms are rattans, e.g. climbing palms. The subtribe Ancistrophyllinae (subfamily Calamoideae) contains most of these with 21 species represented by three genera: Eremospatha, Laccosperma, and Oncocalamus. African rattans are mainly restricted to the tropical rain forests of Central and West Africa and play an important role in the local economy thanks to their use in furniture making. Here, we present a species level phylogeny of the Ancistrophyllinae based on four plastid intergenic spacers (psbA-trnH, psbZ-trnfM, atpI-atpH, and rps3-rpl16) sequenced for 80% of the species found in the subtribe (17/21). Data were analyzed using maximum parsimony, maximum likelihood, and Bayesian inference methods. Our results show that subtribe Ancistrophyllinae is strongly supported as monophyletic, as are the three genera. However, relationships among the genera remain unresolved. Our analyses provide details on the relationships among species within Eremospatha and Laccosperma, but not in Oncocalamus. The species E. cabrae and E. dransfieldii were recovered with strong support as sister, together forming a highly divergent clade sister to the rest of the genus. The four plastid markers used here provide useful levels of resolution and could be considered in other species-level analyses in palms.
The genus Louisiella (Poaceae, Paniceae) is reviewed. This African monotypic genus was originally circumscribed on the basis of morphological characters. Recent phylogenetic studies have shown its affinity with an American species of Panicum, P. elephantipes. Molecular, morphological, and anatomical characters of Louisiella fluitans and P. elephantipes were analyzed. Both species share vegetative and reproductive characters, including habit, ligules, blades, inflorescence, and spikelet features, such as a reduced lower glume, lanceolate upper glume and lower lemma both longer than the upper anthecium, lower palea reduced or absent, and lower flower absent, upper anthecium indurate, and a caryopsis with a linear hilum. Anatomically, both taxa are Kranz of the PS-subtype, with specialized chloroplasts on the outer parenchymatous sheath. The molecular phylogenetic analysis also corroborates the relationship between these species, with a 100% bootstrap support. As a result, we conclude that P. elephantipes has to be transferred to Louisiella, making that genus amphitropical in distribution. Affinities of Louisiella with other genera of Paniceae are also discussed. Finally, two lectotypes are designated for Panicumfistulosum and Digitaria megapotamica.
Multivariate and univariate analyses based on morphological characters were carried out in order to understand the taxonomic status of Senna spiniflora and S. chacoënsis. Fifteen morphological characters from herbarium specimens were recorded and analyzed by the Kruskal-Wallis test and principal components analysis. The type specimens and all available herbarium material were examined for the taxonomic study. The results showed that the taxa differed only by the character of stem pubescence and this feature displayed high variability. In parallel, the principal components analysis demonstrated that no clearly separated groups were recognized. Based upon these results, the synonymy of S. chacoënsis to S. spiniflora is proposed.
Rhynchosia sect. Polytropia is restricted to South Africa and comprises two closely related species (R. ferulaefolia and R. pinnata), which are characterized by pinnately or bipinnately plurijugate, or decompound leaves. The identity of the latter species is, however, uncertain. Several specimens, mostly from the Caledon and neighbouring areas in the Western Cape Province of South Africa, were erroneously assigned to R. pinnata. These specimens belong to a hitherto undescribed species, here named Rhynchosia smithiana. Although this species lacks the distinctive compound leaves typical of section Polytropia, it resembles R. pinnata and R. ferulaefolia in having a spreading, non-twining habit and congested multi-flowered inflorescences. Diagnostic features of the constituent species of section Polytropia, including the newly described species, are reported. A key to the species of this group is also presented, as well as morphological illustrations and distribution maps.
We present a detailed morphological study of three South American species of Ormosia (Leguminosae, Papilionoideae, genistoid clade) that have been confused in herbaria and in the literature. The three species are morphologically linked by the shared possession of relatively few leaflets, small, thin-walled fruits, and relatively small seeds, and they each occur in habitats associated with white sand deposits. Individually diagnostic morphology and non-overlapping geographical distributions support their specific distinction from each other and from other closely related species of the genus. Ormosia costulata is recircumscribed to include only populations in Suriname and French Guiana, while Ormosia trifoliolata, previously treated as a synonym of O. costulata, is lectotypified and re-established as a distinct species ranging across much of the Brazilian Amazon and western Guiana Shield. Material formerly identified as O. costulata from the Atlantic Forest domain in the Brazilian state of Bahia is recognized as a distinct and endangered species, herein described and illustrated as O. lewisii.
Separate and combined Maximum Likelihood and Bayesian analyses of the papilionoid legume genus Poecilanthe sensu lato (Leguminosae) and other Genistoid genera were performed using molecular (nuclear ribosomal and chloroplast matK) and morphological data. The data are in excellent agreement that Poecilanthe is paraphyletic with respect to other genera of the tribe Brongniartieae. Each of the three independent lineages of Poecilanthe sensu lato combines a morphological, geographical, and ecological distinction. They will be ranked at the genus level and Poecilanthe sensu stricto is recircumscribed to encompass the extra-Amazonian species that occur in highly seasonal dry forests.
The South-American genus Amphiodon Huber is reinstated and circumscribed to include a single species, A. effusus. The genus is segregated from Poecilanthe and is diagnosed by a vexillary stamen that is mostly adherent to the standard claw and a septate pod with seeds that are transversally arranged. Poecilanthe ovalifolia is synonymized with A. effusus. Description, illustration, and geographical distribution for A. effusus are provided.
A new Oxalis (Oxalidaceae) species from the Hantamsberg (South Africa) is described and typified as Oxalis noctiflora using morphological, palynological, and molecular characters. Superficially it closely resembles Oxalis aridicola, Oxalis callosa, Oxalis odorata, and Oxalis primuloides, and it shares a unique supra-areolate pollen type with most of these species. No single vegetative morphological character defines this species, but a unique combination of bulb and leaflet characters distinguishes it from all other known species. The flowers of Oxalis noctiflora are unique for the genus in displaying typical moth-pollination characters. The white flowers open at night, have a long, unusually narrow, corolla tube, and produce a strong sweet scent. All previously described Oxalis species have flowers that open during the day. Phylogenetic analyses based on nuclear Internal Transcribed Spacer data placed this species sister to O. aridicola within a clade of morphologically and palynologically similar taxa.
Manihot veadeirensis, from Chapada dos Veadeiros, Goiás, Brazil, is described, illustrated, and assigned to Manihot section Quinquelobae. This new species is similar to Manihot mossamedensis, differing from it in the gamosepalous calyx of its pistillate flowers, lanceolate, persistent, and uninerved bracts and bracteoles, exserted staminate flowers, and wingless capsules. The distribution, habitat, conservation status, and systematic position of the new species are presented. A key to all species of Manihot section Quinquelobae is also provided.
Two new species of Passiflora subgenus Deidamioides section Tetrastylis (Passifloraceae) from Brazil, Passiflora igrapiunensis and P. timboënsis, are described and illustrated. Taxonomic comments on the related species are also included.
A new species of Leandra from the state of Tocantins, Brazil, is described and illustrated here. Leandra heteroporata can be recognized by the slightly anisophyllous leaves covered by simple glandular trichomes (1–6 mm long), two to four lateral and terminal inflorescences per node and the stamens with linear-subulate anthers slightly different in size and shape, which are basally bifurcated in the larger stamens. The morphology and geographical distribution of the new species are compared with its closest related taxa. Leandra heteroporata is the first record of the Leandra sensu stricto clade from the state of Tocantins.
During a floristic survey of Melastomataceae carried out in Biribiri State Park and around Diamantina, Minas Gerais State, Brazil, we found a new species of Microlicia, which is endemic to campos rupestres from the Espinhaço Mountain Range. Microlicialongicalycina has flowers with long calyx lobes, a dense indumentum of glandular trichomes on the hypanthium, leaves with a light green and ciliate margin, and with a mix of glandular trichomes and pale, thickened, long trichomes. A discussion on the leaf anatomy of the species is provided.
Echinocereus is the third most species-rich genus in the Cactaceae. It is distributed in North America from Mexico to the central U. S. A. Previous molecular phylogenetic studies have indicated that the genus is polyphyletic, but incomplete taxon sampling and unclear resolution have hindered the formal re-evaluation of generic and infrageneric circumscriptions. To address this problem, we analyzed six plastid regions (matK, rbcL, psbA-trnH, trnQ-rps16, rpl16, and trnL-F) using maximum parsimony and Bayesian inference criteria for 59 species, including all previously proposed infrageneric entities and representing the full range of morphological variation known in the genus. Our results support the monophyly of Echinocereus if E. pensilis is excluded and reestablished as the monotypic genus Morangaya. Two additional morphological characters, erumpent flower buds and green stigma lobes, further support the circumscription of Echinocereus sensu stricto. Phylogenetic analyses recovered nine main clades in Echinocereus s. s., one of which corresponds to the Triglochidiati section; the remaining clades did not correspond to any other recognized sections. We suggest a re-evaluation of previously proposed infrageneric entities.
Strumpfia maritima is a morphologically unique member of the Rubiaceae that has held several taxonomic placements for a long time. In the most recent tribal delimitations within the subfamily Cinchonoideae, Strumpfia was included in the tribe Chiococceae, based solely on molecular phylogenetic positioning, despite the presence of significant morphological and palynological differences between Strumpfia and the other genera of the tribe. Previous studies refrained from including Strumpfia in the Chiococceae because of its numerous morphological and palynological differences, and the inclusion of Strumpfia creates a tribe without a single synapomorphy. Our study of genetic divergences among the tribes of the subfamily Cinchonoideae using trnL-F data, analyzed using both maximum parsimony and Bayesian inference, provides additional evidence for including Strumpfia in the monotypic tribe Strumpfieae.
Little is known about the sexuality of the woody Neotropical genus Aegiphila. For a long time species of Aegiphila were thought to be heterostylous, but more recently it has been reported that a few species are dioecious and suggested that likely most Aegiphila are functionally dioecious. In this note, dioecy is confirmed in Aegiphila odontophylla from live plants in the Talamanca mountains in Costa Rica. Further, comparison of total flower size revealed that male and female reproductive structures are displayed at similar heights, likely resulting in accurate pollen uptake and delivery by pollinators. The length of the corolla tube as an estimate of flower showiness on the other hand was significantly different between male and female individuals. Male individuals had longer corolla tubes, suggesting a possible role for sexual selection in the evolution of floral dimorphism in this species.
The large and economically important genus Solanum contains ca. 1,400 species distributed worldwide. One of the 12–14 major clades identified in the genus is the Leptostemonum clade, or tire “spiny solanums.” Previous molecular phylogenetic studies have identified 14 major clades in the spiny solanums. Among these is the Carolinense clade, which contains Solanum carolinense, the type species of S. subsect. Lathyrocarpum. However, previous results have suggested that the species composition of the Carolinense clade is only partially congruent with taxonomic circumscriptions of S. subsect. Lathyrocarpum. The main goal of this study was to increase taxon sampling over previous molecular phylogenies in order to better understand the composition of the Carolinense clade. We newly sequenced 31 accessions of 17 taxa putatively belonging to the clade for two nuclear (ITS, waxy) and one plastid gene region (trnT-trnF) and combined these with previously generated molecular data. Phylogenetic analyses of separate and combined DNA matrices were done using maximum parsimony and Bayesian methods. Results from analyses of the combined nuclear and plastid data set showed 10 taxa to be resolved within a well-supported Carolinense clade. However, three species, Solanum dimidiatum, S. perplexum, and S. hieronymi, showed conflicting placements in phylogenies based on analyses of nuclear vs. plastid data sets, suggesting a history of allopolyploidy or introgression. Within the Carolinense clade, the North and South American species were both recovered as well-supported clades, implying a single dispersal event from South to North America.
Plants that have adapted to grow as submersed aquatics are relatively rare among angiosperms, yet they represent a convergent strategy that has evolved in many groups. Asteraceae tribe Eupatorieae include a number of obligate wetland species, but only three genera (Gymnocoronis, Sclerolepis, and Shinnersia) routinely grow submersed. We conducted a molecular phylogenetic study of Eupatorieae to assess the relationships among aquatic genera and to infer the evolutionary history of the submersed aquatic habit. Two major clades were identified in Eupatorieae, corresponding to base chromosome numbers of x = 10 or fewer, and x = 11 or greater. We determined that submersed aquatics evolved independently in two subtribes, Adenostemmatinae (Gymnocoronis; x = 10) and Trichocoroninae (Sclerolepis and Shinnersia; x = 15), each belonging to a separate major clade, and that the aquatic lineages evolved in allopatry. Sparse taxon sampling precluded a firm assessment of ancestral states, although two widespread genera, Adenostemma and Mikania, were implicated as likely relatives of the aquatic lineages. Our data also support the continued recognition of Shinnersia and Trichocoronis, which occasionally have been considered synonymous, as distinct genera.
This taxonomic treatment is the first in the genus Calycera based on a detailed study of morphology and a critical analysis of species boundaries. In this revision, nine species and four varieties are recognized, one new combination (C. crassifolia var. spinulosa), nine new synonymies (Anomocarpus axillaris, Boopis integrifolia, Calycera boopidea, C. crenata, C. foliosa, C. intermedia, C. pulvinata f. cauligera, C. squarrosa, and Gymnocaulus viridiflorus), six new lectotypifications (for Anomocarpus, Boopis gracilis, Calycera crenata, C. pulvinata f. cauligera, C. sessiliflora, and C. sympaganthera), and one neotypification (for Calycera involucrata) are established. Updated morphological descriptions (including an emended description of C. herbacea var. sinuata) and geographical distributions are included for each species. Analyses of their life forms and inflorescence, flower, and fruit morphology are presented.
A new species of Campanula from Central Albania, named C. skanderbegii, belonging to isophyllous Campanula ser. Garganicae is illustrated and described. Morphologically and phylogenetically it is well differentiated from other species of this series. It is morphologically similar to C. portenschlagiana from Central Dalmatia (Croatia) based on corolla shape. Phylogenetic analyses, maximum parsimony, maximum likelihood, and Bayesian inference, based on nuclear ITS and chloroplast trnL—trnF data support C. skanderbegii as a clearly distinct taxon within the isophyllous species of Campanula. A distribution map for C. skanderbegii and a key to all species of Campanula ser. Garganicae are provided. Because of its rarity and highly restricted distribution, C. skanderbegii should be included in the IUCN red list as a critically endangered species.
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