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Rampant homoplasy can be a major challenge in the classification of land plants that have limited morphological differences and/or ecological diversity, such as liverworts and mosses. Here we present the first comprehensive molecular phylogeny of the derived liverwort genus Cololejeunea and its close relatives. We used three markers (trnL—F, rbcL, and nrITS) and 116 accessions representing the geographic range as well as the morphological diversity of this predominantly epiphyllous genus. The molecular data support three major lineages: Colura, Myriocoleopsis, and Cololejeunea. Myriocoleopsis species were resolved in a clade with Cololejeunea minutissima. Aphanolejeunea and Chondriolejeunea were both resolved as putative monophyletic groups, and nested in the Cololejeunea clade. Cololejeunea angustiflora was found to be sister to the clade comprising the remaining Cololejeunea species. Currently accepted subgenera of Cololejeunea were paraphyletic or polyphyletic. Six out of 30 species with multiple accessions were not supported as natural taxa. The current classifications of Cololejeunea have been influenced by frequent homoplasy of morphological characters and do not accurately reflect species relationships. A new classification for this genus is outlined. Our data did not provide evidence for ancient vicariance events where major clades and continents are associated, instead some evidence was recovered for recent intercontinental species range expansion.
The chromosome number and phylogenetic position of the monotypic eusporangiate genus Mankyua (Ophioglossaceae), which shares peculiar morphological characteristics with Ophioglossum s.l. and Helminthostachys, were determined. Meiotic and mitotic chromosome observations of Mankyua show n = 130 and 2n = ca. 260, suggesting that the base number of this genus is x = 130, which is a new finding and the highest base number reported so far for Ophioglossaceae. The molecular phylogenetic trees derived from nucleotide sequence variation of rbcL and matK genes differ in terms of the phylogenetic position of Mankyua. The molecular phylogenetic trees based on combined data sets of rbcL and matK suggest that Mankyua might be the earliest-diverging lineage of Ophioglossaceae.
Two new species of Peperomia from Brazil are described and illustrated based on morphological traits: Peperomia diamantinensis from the state of Bahia and Peperomia cruzeirensis from the state of Acre. Peperomia diamantinensis is very similar to Peperomia obtusifolia and Peperomia magnoliifolia, differing from the latter in the acute leaf apex and the short internodes. Peperomia cruzeirensis is similar to Peperomia elongata Kunth, but differs from the latter in the short pedicel of the fruits, and the petiole without two rows of trichomes. Discussions on taxonomy, geographic distribution, conservation status and habitat are provided, as well as identification keys to the Brazilian species of the Oxyrhynchum and Leptorhynchum clades.
Anthurium is a strictly Neotropical genus of Araceae ranging from southern Mexico to northern Argentina, including approximately 900 accepted species names. Despite its immense diversity, its ecological importance in Neotropical forests, and a long history of botanical collection, cultivation, and taxonomical research, Anthurium has been only cursorily sampled in previous molecular phytogenies. This study aims to test the monophyly of Anthurium, to understand the evolutionary history of the genus, and to elucidate relationships among its species using maximum parsimony, maximum likelihood and Bayesian analyses on a combined chloroplast (trnG intron, tmH-psbA and tmC-ycf6 intergenic spacers) and nuclear (CHS first intron and partial flanking coding regions) DNA sequence dataset for 102 Anthurium species and closely related outgroups. Despite some limitations (ca. 11% species richness coverage, and lack of nuclear sequences for outgroups), results indicate that Anthurium is a strongly supported monophyletic genus and that at least 18 major supported clades are recognizable within it, most of them easily characterized morphologically and/or geographically. This study also suggests that the current sectional classification of Anthurium does not accurately represent its evolutionary history since most of the major clades recovered in these analyses do not correspond with the current circumscriptions of infrageneric groups. Despite using the most variable gene regions available, low sequence divergence was found among Anthurium species, relatively short branches characterize the core of the Anthurium clade, and resolution is still lacking in the deeper nodes of the phylogeny, a pattern consistent with a rapid, and probably recent, radiation of species.
A phylogenetic analysis of the genus Homalomena (Araceae: Homalomeneae) based on the nITS region is presented. Eighty-nine taxa are included; representing all Asian supergroups, several Neotropical species currently assigned to Homalomena, and selected species of Philodendron. Asian Homalomena is well supported as monophyletic and excludes Neotropical Homalomena. The Cyrtocladon supergroup is monophyletic after transferring the Insignis complex and Havilandii complex into the Punctulata supergroup. The Homalomena and Chamaecladon supergroups are well supported. A reduced phylogeny of 20 accessions representing 15 taxa was used for subsequent morphological and chemical marker optimization. A constricted spathe and four stamens per staminate flower are plesiomorphic for Homalomena. Staminodes among the pistillate zone are lost three times independently in Homalomena supergroup, Punctulata supergroup, and H. vivens. Chemical markers identified from liquid chromatography-mass spectroscopy profiling provided an independent set of markers that further support the separation of Neotropical species of Homalomena from the Asian taxon. Three chemical markers at Rt 2.55 min, 2.69 min, and 2.90 min are shared among the majority of taxa sampled for Asian Homalomena, and Neotropical species currently assigned to Homalomena show two unique peaks at Rt 3.25 min and 3.54 min. Five chemical markers support the Cyrtocladon supergroup with the exception of Homalomena vivens. A chemical marker at Rt 3.60 min is plesiomorphic for the Chamaecladon, Homalomena and Punctulata supergroups. A chemical marker at Rt 2.80 min is apomorphic for the Chamaecladon supergroup, with a separate gain in H. punctulata. This study supports removal of the Neotropical species from Homalomena, redefines the morphological boundaries of Homalomena sensu stricto (i.e. the Asian species), and supports and refines the grouping of Asian species into supergroups.
Borneo is one of the richest areas for mycoheterotrophic plants, and six species including one endemic have been recorded for the genus Sciaphila Blume, Triuridaceae. Here, we recognize two previously undescribed species of Sciaphila from botanical surveys in Betung Kerihun National Park, West Kalimantan, Borneo, and provide detailed morphological accounts of both new species, S. betung-kerihunensis and S. brevistyla. We also provide a key to the species of Sciaphila in Borneo.
A revision of the Cohniella cebolleta complex from northern South America and the Lesser Antilles is presented. Five species are recognized in this complex. As circumscribed here, Cohniella cebolleta is restricted to northern Colombia and northern Venezuela. We describe and illustrate a new species, Cohniella macrocebolleta, endemic to eastern Venezuela, and propose two new combinations: Cohniella juncifolia, endemic to the Lesser Antilles, and Cohniella ultrajectina from southern Venezuela and the Guianas (Guyana, Suriname, and French Guiana). Detailed taxonomic descriptions are given for each species, including short accounts of their distribution and ecology, diagnostic characters, variational range, taxonomic comments, and material examined. Furthermore, a key, an appendix featuring diagnostic characters, and distribution maps for species complexes recognized in Cohniella and taxa recognized in the C. cebolleta complex are presented and discussed.
A new grass species from Brazil, Paspalum giuliettiae, is herein described, illustrated, and compared with some congeners. This new species shares inflorescences with subconjugate primary branches and solitary glabrous spikelets with P. pumilum, but it differs in macromorphological characters and in leaf anatomy. Paspalum giuliettiae has been collected in the Chapada Diamantina, which is the northern portion of the Espinhaço Range, in the central region of the State of Bahia, Brazil. This new Paspalum grows in humid or swampy places in the ‘campos rupestres’ vegetation, above 1,000 m, and seems to be endemic to this region.
In advance of a comprehensive systematic study of New World Mucuna, we provide a taxonomic revision of the species of Mucuna that occur in Brazil. A new species, Mucuna analucianae, endemic to eastern and central Brazil, is described and illustrated. Mucuna huberi is designated as a synonym of M. elliptica, and a lectotype for M. elliptica is chosen. Mucuna pluricostata is considered to be a synonym of M. pruriens, and a lectotype for M. pluricostata is chosen. Mucuna eriocarpa is excluded from the genus. We recognize seven species of Mucuna in Brazil, only two of which are endemic to the country. Two of the seven species are assessed as endangered (EN) according to IUCN criteria.
Crotalaria sect. Amphitrichae is described as new to give formal recognition to a strongly supported clade revealed by a recent molecular systematic study. The five known species are restricted to the arid western parts of southern Africa (C. colorata, C. excisa, C. humilis, C. meyeriana, and C. pearsonii). Species of the section are morphologically distinct in having (1) hairs distributed all around the distal part of the style; (2) ridge callosities on the standard petal blade and claw; (3) a rounded keel; (4) a twisted keel beak; and (5) a rounded style. A cladistic analysis of 18 morphological characters resulted in a well-resolved cladogram that recovers relationships within the section. The study revealed that two new species should be recognised, described here as C. giessii and C. kolbergii. A taxonomic revision of the section is presented, with descriptions, diagnostic characters, distribution maps, and illustrations of all seven recognised species.
The present taxonomic revision of Ficus subsection Urostigma recognizes 27 species, of which three are new: F. chiangraiensis, F. middletonii, F. pseudoconcinna. Two new varieties are distinguished within F. virens, var. dispersa and var. matthewii. Ficus lecardii and L. salicifolia, formerly subspecies of F. cordata, are again reinstated to the species level. Typical characters for the subsection are monoecy, monostaminate flowers, red(-brown) colored ovaries and cystoliths on only the abaxial leaf surface. Ficus amplissima and F. rumphii (section Leucogyne) were formerly part of subsection Urostigma, and they have been added here to the key and descriptions because of their morphological resemblance with the species in F. subsection Urostigma. Molecular based phylogenetic analyses showed that at least F. rumphii is unrelated to subsection Urostigma. The two species only differ from subsect. Urostigma in their whitish ovaries and cystoliths at both sides of the leaf blade and they are pollinated by a different group of wasps, species of Eupristina subg. Parapristina.
Sorocea longipedicellata (Moraceae) is described and illustrated as a species new to the Atlantic rainforest domain. The new species is related to the group of Sorocea species with completely spinulose dentate leaf margins. It includes Sorocea bonplandii and S. guilleminiana, but is distinguished mainly by elongated pedicels when fruiting. This species is known only from Bahia state, northeastern Brazil and is designated as a critically endangered species because its extent of occurrence is estimated to be less than 100 km2 in an extremely fragmented area.
Phylogenetic relationships of Passiflora subgenus Decaloba were examined using 148 taxa and four molecular markers: nuclear nrITS, ncpGS, cp trnL-F, and ndhF. Relationships of subgenus Decaloba to the other four Passiflora subgenera (Astrophea, Deidamioides, Tetrapathea, and Passiflora) are investigated, as are relationships among the eight supersections within subgenus Decaloba. Results indicate that subgenus Deidamioides is not monophyletic. Subgenus Astrophea subgenus Deidamioides (section Tryphostemmatoides) together form the most basally branching lineage in the genus, followed by a clade comprised of subgenus Passiflora subgenus Deidamioides (sections Tetrastylis, Polyanthea, and Deidamioides). Passiflora obovata (subgenus Deidamioides section Mayapathanthus) is resolved as part of subgenus Decaloba. The Old World subgenus Tetrapathea is supported as sister to subgenus Decaloba. Subgenus Decaloba is monophyletic and contains seven major lineages that generally correspond to currently recognized supersections. Within subgenus Decaloba, supersection Pterosperma is most basally branching, followed by supersection HahniopathanthusP. obovata. The New World species Passiflora multiflora, the type of supersection Multiflora, is resolved as sister to a monophyletic Old World supersection Disemma. The remainder of the former supersection Multiflora is paraphyletic with respect to supersection Auriculata. Supersections Cieca, Bryonioides, and Decaloba are monophyletic. Within supersection Decaloba, two main clades are resolved: 1) section Xerogona section Decaloba pro parte and 2) the remainder of section Decaloba. The molecular phylogeny supports a New World origin for Passiflora, with two independent radiations to the Old World. Morphological synapomorphies are discussed for the major clades, documenting a pattern of remarkable evolutionary lability in several notable characters.
Two new species of Marcetia (Marcetia paganuccii and M. cardosoana) are described and compared with their closely related species. The two new species were found exclusively in mountainous areas of the Chapada Diamantina, Bahia, Brazil. Illustrations, distribution maps, conservation status and SEM photographs of leaves and seeds for both species are provided.
The present study aimed to develop a robust, extensively sampled phylogenetic hypothesis across all species in Leavenworthia and Selenia using chloroplast (psbJ—petA) and nuclear encoded (nrDNA ITS and LUMINIDEPENDENS) DNA sequence data. Parsimony and maximum likelihood methodologies were employed. Strong maximum likelihood bootstrap and parsimony jackknife support were recovered for the monophyly of the sister pair Leavenworthia plus Selenia and for the respective monophyly of each genus. Leavenworthia and Selenia are hypothesized to have diverged in the Ozark Mountains and results from an analysis of branch lengths leading to Leavenworthia and Selenia clades are consistent with Leavenworthia having diversified more recently than Selenia. Polyploidy and extensive aneuploidy complicate the interpretation of chromosomal evolution within and between these genera. Comparative results from phylogenetic analyses indicate that L. aurea and L. texana are hybrids with a shared set of putative parents. Lastly, patterns of diversification within Leavenworthia appear to have included a minimum of three transformations from self-incompatible to self-compatible systems.
Allopolyploidy is common in angiosperms, but only rarely involves different genera. One hypothesized case of intergeneric allopolyploidy is Hylocereus megalanthus, a member of Cactaceae tribe Hylocereeae, a group of vine cactus species, some of which are known for their edible fruits (“pitahaya” or “dragon fruit”). This polyploid species has been interpreted as morphologically intermediate between Hylocereus and Selenicereus. Plastid and nuclear ribosomal DNA ITS sequences from all H. megalanthus individuals sampled are either identical (plastid), or form a monophyletic clade despite considerable intraindividual polymorphism (ITS). Plastid and ITS phylogenies both show H. megalanthus nested within a well-supported Hylocereus, which in turn is nested within a paraphyletic Selenicereus. The absence of more than one lineage of ITS in H. megalanthus is consistent with autopolyploidy, but could be due to inter-homoeologue concerted evolution. Numerous low-copy nuclear genes were tested for utility in these vine cacti, and two (phyC and Vatp1) were sampled from H. megalanthus and a subset of Hylocereus and Selenicereus species. In both cases, H. megalanthus haplotypes were more closely related to each other than to other Hylocereus or Selenicereus haplotypes. Thus, we found no evidence for allopolyploidy, let alone intergeneric allopolyploidy, in H. megalanthus.
The tribe Phyllodoceae is comprised of seven genera (Bejaria, Elliottia, Epigaea, Kalmia, Kalmiopsis, Phyllodoce, and Rhodothamnus). A clade within this tribe, comprised of all genera except Bejaria, was the subject of a phylogenetic study using six molecular markers. The goal of the study was to reconstruct a species-level phylogeny for the purpose of examining the evolution of morphological characters and to provide a framework for future studies in historical biogeography. Twenty-nine of 31 Phyllodoceae species were included in the molecular dataset of sequences from four plastid and two nuclear regions, and the remaining two species were included in a small morphological analysis. Most nodes were strongly supported using combined molecular data and four analytical approaches. Potential morphological synapomorphies were identified for some clades within the group. Evolutionary patterns in some morphological characters are apparent, which provide a basis for future anatomical and developmental studies.
This paper provides descriptions and illustrations of a new species, Barjonia furlanii (Asclepiadoideae, Apocynaceae), from Serra da Canastra National Park, in the state of Minas Gerais, Brazil. This species is similar to B. laxa, especially in its frondose thyreoid inflorescence, compound corona, and linear leaves. It is, however, distinguished from B. laxa by internodes (up to 4 cm long), peduncles (4.2–5mm long) and much shorter pedicels (4–6 mm long), erect leaves, attenuated at the base, and partial inflorescences that alternate with 2–4 flowers. B. furlanii is proposed for Endangered status under IUCN guidelines (EN B2abiii), and a key to all Barjonia species is included in this study.
The new species Borreria krapocarmeniana (Rubiaceae) is described and illustrated. The taxonomic identity of Borreria linoides and Borreria scabiosoides are discussed. Borreria organensis and Borreria rubricaulis are proposed as new synonyms of B. scabiosoides. Seed micromorphology and pollen grains of the three species are analyzed and a map with the distribution of these species is presented. Additionally, a table featuring diagnostic characters to distinguish between species is provided.
Pogogyne (Lamiaceae) is a small genus of annual plants, occurring almost entirely in vernal pool habitats. To infer the phylogenetic relationships of this group, DNA sequence data from the trnQ-rps16 chloroplast spacer and both ETS and ITS nuclear regions were obtained from all seven extant species and several outgroups and analyzed using both parsimony and Bayesian phylogeny reconstruction methods. Ancestral state reconstructions were performed for morphological characters that have been used to separate groups within Pogogyne. This study indicates that subgenus Hedeomoides is monophyletic with good support and can be diagnosed by two apomorphies: a reduction in stamen number and a reduced corolla size. Subgenus Pogogyne is paraphyletic, but with poor support. Several clades within the genus Pogogyne have discrete biogeographic distributions. The Pogogyne genus clade has a relatively long stem lineage, but its crown clade has quite short branches. Our estimate for the mean divergence time of the stem node is 5.1–7.7 million years ago and for that of the crown node is 0.9–1.9 million years ago, the latter overlapping with ages of vernal pool ecosystems. These results support the hypothesis that the members of Pogogyne underwent a rapid diversification in response to specialization to a periodically inundated habitat.
Members of the small genus Conopholis are perennial holoparasites. They are found growing in eastern and southwestern North America and in Central America, where they attach to the roots of their oak hosts. Two species were recognized in the last taxonomic revision of the group based on geographic range and differences in floral, capsule, and bract morphology. Due to the overlapping nature of the characters used to distinguish between taxa, no single morphological feature can be relied on to differentiate between the species. A recent molecular phylogenetic study of the genus recovered three well-supported lineages, none of which corresponds entirely to the current subdivision of the genus into two species. We undertook a fine-scale morphometric study of the genus, emphasizing calyx and bract morphology. Unweighted pair-group method using arithmetic averages and principal coordinate analyses corroborate molecular data and strongly support the distinction of three separate lineages within Conopholis. A taxonomic re-alignment is proposed for the genus including three species, C. americana, C. panamensis, and C. alpina, each with various degrees of overlap with previously described taxa.
In spite of the recent studies on the phylogeny of Verbenaceae, the position of the monotypic Verbenoxylum, endemic to the Atlantic rainforest in southeastern Brazil, remains unsolved. Molecular data were here analyzed to infer the phylogenetic placement of this genus; furthermore morphological data was studied in order to examine traits that support relationships among taxa. Sequences of the plastid regions of ndhF gene and trnL—trnF intergenic spacer were analyzed to conduct phylogenetic studies with maximum parsimony, maximum likelihood, and Bayesian inference. Morphological traits that had been traditionally used to distinguish tribes within Verbenaceae, as well as those employed to characterize Verbenoxylum, were examined. Verbenoxylum is nested within the tribe Duranteae, sister to Recordia, a monotypic genus endemic to Bolivia, a placement never reported before. The morphological traits analyzed prove not to be useful to distinguish tribes but are important at lower taxonomic levels. Based on the sister relationship and morphological similarities between the genera Verbenoxylum and Recordia, we propose the inclusion of Verbenoxylum reitzii into Recordia, forming the new combination Recordia reitzii.
Phylogenetic analyses were performed based on four plastid DNA sequences to determine if the Daturae and Datura are monophyletic and to understand their relationships and classification. Species of Datura have been divided into three sections based upon the position and type of the fruit: Ceratocaulis, Datura, and Dutra. Our analyses confirmed the monophyly of the Daturae clade with two genera, Datura with herbaceous species and Brugmansia with woody species. The Datura clade consists of two distinct lineages, that of D. ceratocaula, corresponding to the monospecific section Ceratocaulis, and one that gives rise to the remaining species of the genus. The large clade readily divides into two groups but they do not correspond to the conventional classification of the genus. Species with regularly dehiscent capsular fruits are present in both groups while species with irregularly dehiscent capsules are found in only one group. Because section Dutra is polyphyletic, realignment of species within two traditional sections is necessary as follows: Datura containing D. arenicola, D. discolor, D. ferox, D. quercifolia, D. kymatocarpa, D. leichhardtii, and D. stramonium; Dutra containing D. inoxia, D. lanosa, D. metel, D. reburra, and D. wrightii. The seed margin clearly distinguishes these sections, rather than the combination of fruit position and dehiscence which has been used previously.
A revision of the genus Stomatanthes (Asteraceae, Eupatorieae) is presented. A cladistic analysis of 43 qualitative and eight quantitative morphological characters resulted in one tree that suggests a narrower delimitation of the genus relative to previous circumscriptions. Stomatanthes thus comprises three species, one widespread in sub-Saharan Africa and the other two endemic to restricted areas in central Africa. A cluster analysis and a non-metric multidimensional scaling were also performed. Of the 25 quantitative traits originally measured for the analysis, eight (blade length and width, involucre length, corolla length, corolla limb width, corolla tube width, thecae length, anther apical appendage length) provided clustering information and indicated the same groupings as the cladistic analysis. This revision also includes full descriptions of species, photographs, illustrations, and distribution maps.
We studied the phylogeny of 22 accessions of Daucus and seven accessions in related genera, with DNA sequences from eight nuclear orthologs and one plastid (psbA/trnH) region. Maximum parsimony and maximum likelihood analyses of the concatenated data matrix of 7,212 aligned nucleotides provided excellent bootstrap support for many clades. Concordant with prior molecular results Pseudorlaya pumila was firmly imbedded within Daucus, as was Margottia gummifera, a new finding. All accessions of D. capillifolius, D. carota, and D. sahariensis formed a 2n = 18 clade with all other species within the Daucus clade with chromosome numbers of = 20, 22, and 44 (D. glochidiatus). Sister to the D. carota clade was a clade containing Margottia gummifera and Pseudorlaya pumila, sister to these species was D. crinitus, sister to all the above was D. muricatus, and sister to all of the above was a clade containing the remaining Daucus species. Bayesian analyses of individual regions analyzed separately and averaged over multiple trees with *BEAST software (a coalescent approach), however, provided a phylogeny at variance with the concatenated approach, most notably in firmly imbedding Turgenia latifolia within Daucus.
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