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Based on morphological and molecular evidence, a new combination, Syntrichia kingii, is proposed for Tortula kingii, an Andean taxon previously treated as conspecific with S. percarnosa. Tortula nigra is also synonymized with S. kingii. Maximum parsimony and Bayesian analyses of chloroplast DNA sequences (trnL-trnF and trnG regions) were undertaken to assess the affinities between these cucullate-leaved species. Syntrichia kingii is described, illustrated, and its distribution mapped. The known range of this species is extended in the Neotropics to Guatemala, Mexico, and Venezuela. Diagnostic characters, microphotographs, and a morphological comparison with S. percarnosa are provided. Syntrichia percarnosa is excluded from the Mexican moss flora.
Dicranum is a large (ca. 90 species) and taxonomically complex moss genus. Circumscriptions and relationships of many Dicranum species remain ambiguous due to the absence of a worldwide revision and comprehensive phylogenetic analyses. In this study, we address species circumscriptions and relationships of presumed close allies within Dicranum sect. Dicranum. Molecular phylogenetic reconstructions based on five chloroplast regions and nrITS suggest a close relationship between D. bonjeanii, D. howellii, D. nipponense, D. japonicum, D. cf. lorifolium, and D. scoparium, which can be regarded as the D. scoparium species complex. In contrast, D. majus and D. polysetum, as well as D. fuscescens and D. spadiceum (former varieties of D. scoparium), are separated from the complex. Molecular data are generally congruent with the morphological species concept, but the circumscriptions of D. bonjeanii, D. japonicum, D. cf. lorifolium, and D. scoparium need further study. Most analyzed D. scoparium specimens from across its Holarctic distribution are contained in one clade (D. scoparium s. s.), but a number of North American specimens are resolved as closely related to D. japonicum and D. cf. lorifolium. Costa cross sections and characters of the leaf apex (shape, serrulation of margins) are most relevant for identifying the studied Dicranum species morphologically.
Adiantum alan-smithii (Pteridaceae) is described from Chiapas, Mexico. It is similar to A. raddianum, and previously identified as such, however it can be distinguished by its yellow-farinose indusia, the proximal acroscopic pinnule of each pinna usually overlapping the main rachis, flabellate segments, and rachises that are weakly flexuose distally. In contrast, A. raddianum has non-farinose indusia, the proximal pinnule of each pinna does not overlap the main rachis, and segments are obovate to rhomboid. The new species is described, illustrated, and a key to species of the A. raddianum group in Mexico is provided. This contribution adds an additional endemic species of Adiantum to the flora of Mexico, and is part of our ongoing effort to improve the circumscription of A. raddianum, one of the most over-applied names in the Neotropical fernflora.
Tectaria brauniana and T. nicotianifolia are unusual in their genus by having creeping rhizomes and two-ranked leaves. Tectaria brauniana is further unusual by having free veins. These rhizome characters of both species, and the free veins of T. brauniana, are typical of Triplophyllum and suggest a relationship to that genus instead of Tectaria. To determine the phylogenetic relationships of T. brauniana and T. nicotianifolia, we used molecular evidence from four plastid DNA markers (rbcL, rps4-trnS, trnG-trnR, and trnL-trnF) to generate a phylogenetic hypothesis for the Tectariaceae. The analysis included the tectarioid genera Arthropteris, Hypoderris, Psammiosorus, Pteridrys, Tectaria, and Triplophyllum. Tectaria brauniana and T. nicotianifolia were recovered as sister to Hypoderris brownii, and these three species were sister to Triplophyllum. These two clades were sister to the rest of Tectaria. Thus, to preserve the monophyly of Tectaria, T. brauniana and T. nicotianifolia are here classified in Hypoderris, a genus previously considered monotypic. We make the following new combination: H. nicotianifolia. In this expanded sense, Hypoderris is characterized by creeping rhizomes, two-ranked leaves, and spiny perispores. The genus occurs in the Caribbean region, Central America, and the Andes from northern Venezuela to Bolivia. A key and illustrations are given for the three species now recognized in Hypoderris.
Meiogyne (Annonaceae) currently comprises 15 species of trees and shrubs, distributed in India, Southeast Asia, Australasia, Micronesia and Polynesia. Previous molecular phylogenetic studies have shown that the Australian endemic genus Fitzalania (consisting of only two species) is nested within Meiogyne, and preliminary morphological data have indicated that several south Pacific Polyalthia species may be misclassified and also associated with Meiogyne. We use maximum parsimony, maximum likelihood and Bayesian analyses based on seven chloroplast regions (matK, ndhF, ndhF-rpl32, rbcL, rpl32-trnL, trnL-F and ycf1) to reconstruct the most comprehensive phylogeny of Meiogyne available to date, inclusive of 13 Meiogyne species, the two Fitzalania species, as well as four Polyalthia species from Fiji and Tonga. The results show that Fitzalania and the four Polyalthia species are nested within Meiogyne, and that two species, M. cylindrocarpa and M. stenopetala, are not natural as currently defined. Meiogyne cylindrocarpa subsp. trichocarpa and M. stenopetala subsp. insularis are not conspecific with their respective autonymic subspecies and are morphologically distinct. Based on the plastid marker phylogeny, and corroborated by morphological observations, both subspecies are elevated to species rank, and the four Polyalthia species and the two Fitzalania species are transferred to Meiogyne, thereby increasing the number of species in the genus to 24.
Persea pumila, a new species of Lauraceae from a Brazilian high-altitude grassland is described and illustrated. The new species belongs to Persea subg. Eriodaphne. It is distinguished from other neotropical Persea species by the shrubby habit in combination with obovate, glabrous, abaxially papillose leaves densely overlapping each other, glabrous inflorescences, and sparsely pubescent flowers. Its morphological distinctiveness and its relationships with probable allied species are discussed.
The monotypic genus Zigadenus occupies a critical position in the evolution of Melanthieae as sister to the clade comprising the rest of the genera in the tribe. Meiotic (n = 27) and mitotic (2n = 54) chromosome counts for Zigadenus glaberrimus documented here do not support a long-standing tentative report of 2n = 52. The likely base chromosome number of the tribe and significance of chromosome numbers as generic synapomorphies are discussed in reference to these newly recorded counts for Zigadenus glaberrimus, a likelyhexaploid.
The lily family Colchicaceae consists of geophytic herbs distributed on all continents except the Neotropics. It is particularly diverse in southern Africa, where 80 of the 270 species occur. Colchicaceae exhibit a wide range of ploidy levels, from 2n = 14 to 2n = 216. To understand where and how this cytogenetic diversity arose, we generated multilocus phylogenies of the Colchicaceae and the Colchicum clade that respectively included 85 or 137 species plus relevant outgroups. To infer the kinds of events that could explain the observed numbers in the living species (dysploidy, polyploidization, or demi-duplication, i.e. fusion of gametes of different ploidy), we compared a series of likelihood models on phylograms, penalized likelihood ultrametric trees, and relaxed clock chronograms that contained the 58 or 112 species with published chromosome counts. While such models involve simplification and cannot address the processes behind chromosomal rearrangements, they can help frame questions about the direction of change in chromosome numbers in well-sampled groups. The results suggest that dysploidy played a large role in the Colchicaceae, with the exception of Colchicum itself for which we inferred frequent demi-duplication. While it is known that triploids facilitate the fixation of tetraploidy and that plant species often include individuals of odd ploidy level (triploids, pentaploids), we hesitate to accept the phylogenetically inferred scenario without molecular-cytogenetic work and data from experimental hybridizations.
We show strong geographical patterning in the morphological and especially the anatomical variation in the widespread southern African danthonioid grass species, Tenaxia disticha. Four varieties can be discriminated, and are formally recognized here as Tenaxia disticha var. disticha, var. vlokii H. P. Linder, var. dracomontana H. P. Linder and var. lustricola H. P. Linder. Tenaxia disticha var. disticha is widespread in eastern southern Africa, from Cape Agulhas at the southern tip of Africa to Inyangani in Zimbabwe; the other three varieties are highly localized: var. vlokii in the southern Cape and vars. dracomontana and lustricola on the high plateau of the Drakensberg. Plastid DNA nucleotide variation revealed only limited geographical pattern, but all regions and all varieties have numerous haplotypes, with no haplotype-poor areas. This is consistent with a model wherein this grass species has occupied its current distribution range for a long time, possibly since the Late Miocene, thus allowing the accumulation of local haplotype diversity. The divergence of the Drakensberg varieties may have been stimulated by the Pliocene uplift of these mountains resulting in a sub-alpine climate. This phylogeographical model is consistent with the hypothesis that southern Africa was not subjected to the dramatic Pleistocene climatic fluctuations driving extensive range changes which were typical of the temperate Northern Hemisphere phylogeographical patterns.
Rhodiola (Crassulaceae) comprises about 70 species and shows a high level of morphological diversity. The genus is mainly distributed in alpine areas and cold regions of the Northern Hemisphere with the Qinghai-Tibetan Plateau as its center of diversity. The phylogenetic relationships within Rhodiola remain poorly understood largely because of difficulties in collecting samples and specimens. In the present study, 51 species/varieties from Rhodiola representing all morphological sections were analyzed using sequences of the nuclear ribosomal internal transcribed spacers region and the plastid psbA-trnH and trnL-F markers. Our results suggest that Rhodiola and Pseudosedum form a well-supported clade, but the relationship between these two genera is not resolved. None of the four currently recognized subgenera is monophyletic; yet three of seven sections, R. sects. Trifida, Prainia, and Pseudorhodiola are each supported to be monophyletic. Rhodiola rosea is a popular medicinal plant that has an adaptogenic effect. The three accessions of R. rosea from eastern Asia, eastern North America, and Europe form a well-supported clade. Rhodiola species independently reached North America from Asia twice, once in the R. rosea lineage, and the other in the R. integrifolia-R. rhodantha lineage. Two taxonomically important characters, dioecy and marcescent flowering stems, are inferred to have evolved multiple times within Rhodiola.
A taxonomic treatment of species of Senegalia (Leguminosae, Mimosoideae) from the Atlantic Domain is presented. We recognized the occurrence of 28 species, from about 52 species in Brazil. The species from the Atlantic Domain occur mainly in these physiognomies: Rain Forest (22), Restinga (14) and Semi-deciduous Forest (12). Fifteen species have wide distributions and 11 taxa have restricted areas of occurrence. We developed a dichotomous key, morphological descriptions, illustrations, and distribution maps for the species. The principal diagnostic characters were the shape, number and position of extrafloral nectaries, number of pairs of pinnae per leaf, and inflorescence type. Two new species are described here: S. amorimii and S. grazielae, and three new records were found for the Atlantic Domain: S. kallunkiae, S. limae, and S. santosii.
A new species of Celastraceae, Maytenus megalocarpa, is described and illustrated. The species is known from only a small number of collections from eastern Brazil, in the Atlantic Rainforest of Bahia and Minas Gerais states. It is considered endangered given the rapid deforestation of this biome. The species is readily distinguished from its congeners by its large, tardily dehiscent fruit that does not open by reflexed valves and by the seeds enveloped by thick arils. Given its remarkable fruits and the consequent initial doubt about the generic position of the new species in Celastraceae, a phylogenetic analysis using molecular data from plastid and nuclear rDNA gene regions, together with morphological characters, was performed. The new species was consistently resolved as nested within Maytenus, which itself is paraphyletic. Incongruence between the nuclear and plastid gene regions was identified for Fraunhofera, Maytenus texana, and Plenckia. Future taxonomic changes are necessary to rigorously delimit Maytenus as a natural genus. Additional comments on geographical distribution, phenology and conservation status of the new species are also provided, as well as a key to the species of Neotropical Maytenus with spinoseleaves.
We describe two new species of Manihot (Euphorbiaceae), and present an updated description of M. pusilla and a synonymization of M. stipularis to M. pusilla. Manihot bellidifolia, and M. longiracemosa found in campos rupestres and cerrado vegetations, reinforce the preference of this genus for habitats with restricted water availability. The two species described here are threatened with extinction, in the category of vulnerable. Manihot stipularis is considered a synonym of M. pusilla due to the wide morphological variation of its stipules, the principal character used to distinguish these species.
Grossera (Euphorbiaceae) is a genus with eight species of trees and shrubs, seven of them from West and Central Tropical Africa and one from Madagascar. It occurs in a wide range of habitats, from primary to degraded forest, and from sea level to 1,350 m. Grossera is characterized by unisexual flowers, terminal inflorescences, anthers inserted on an androphore surrounded by five fleshy glands, three styles divided into two branches, ovary globose inserted in a plane disc thickened at the edge, and fruits in capsule with three 2-valved cocci. Although some regional treatments and a synopsis are available, a comprehensive monograph of the genus is lacking. In addition to a comprehensive key and descriptions, illustrations of five species and distribution maps for all species are provided here for the first time. Taxonomic discussions of each species have been included and three lectotypes are designated. Grossera major, synonymized by some authors, is clearly distinguished from G. paniculata because of its longer pedicels on male and female flowers. The eight accepted species (G. elongata, G. glomeratospicata, G. macrantha, G. major, G. multinervis, G. paniculata, G. perrieri, and G. vignei) are divided into three sections, including a new combination, Grossera sect. Columnella.
Although Stillingia loranthacea has commonly been considered a synonym of Stillingia saxatilis, it is reinstated to specific status in the present study. Stillingia loranthacea differs from S. saxatilis due to its orange-brown to dark brown bark, leaves sessile to subsessile (up to 4 mm long), obovate to elliptic-obovate with apex obtuse to rounded, base attenuate to decurrent, venation brochidodromous, and glands scutelliform. Both species are described and illustrated herein, including observations on habitat and conservation status, a map of distribution, a key, and a table containing distinctive characters. A lectotype is chosen for S. loranthacea.
A new species of Passiflora from the rocky fields (“campos rupestres”) of Minas Gerais State is described: P. rupestris, which belongs to Passiflrora sect. Decaloba. The species is distinguished by having a 3-lobed obtriangular leaf blade that is longer than wide, of small size (1.5–2.9 × 1.3–2.2 cm), without ocelli, and with slightly divergent lobes (27–40°), and a particular combination of characteristics of the flower and fruit. The species is illustrated and its affinities with other species are discussed. Passiflora rupestris must be regarded as critically endangered because of its extremely limited occurrence.
Oenothera, the evening primrose genus, is a model system for studying the evolution of flowering plant reproductive biology. Members of this group vary in the species of pollinator that visit their flowers and in breeding systems, including both self-compatible (SC) and self-incompatible (SI) species. Here, we examine the evolutionary relationships among the six species of Oenothera section Kneiffia using sequences from two nuclear and four chloroplast genes. Through field studies we describe the effective pollinators for four species that had not been previously reported, and experimentally test for pollen limitation. Three of the six species are SC, and three are SI. The phylogeny strongly supports three separate transitions from SI to SC. Despite the expectation that SC species evolved the ability to self because of pollen limitation, or that pollen limitation evolves among SC species, we found no significant differences in pollen limitation between the SI and SC species in this study. Our results resolve the interspecific relationships within section Kneiffia, show that breeding systems can be quite labile, and provide evidence that transitions to self-compatibility do not always coincide with pollenlimitation.
The new genus Quipuanthus from the foothills of the eastern Andes of Ecuador and Peru is described. Quipuanthus seems to be related to Allomaieta, Alloneuron, and Wurdastom in the tribe Cyphostyleae, but the combination of characters as an herb with haplostemonous flowers, recurved style, inferior ovary and apically dehiscent capsular fruits is unique among the Melastomataceae. The new species Quipuanthus epipetricus is described and illustrated.
Dombeya section Hilsenbergia includes five species endemic to Madagascar and the Comoros that are characterized by pendulous inflorescences and long staminal tubes. Morphological data and previous molecular work support the inclusion of these species within Dombeya. Study of morphological characters previously used in the taxonomy of section Hilsenbergia found overlap in these characters among species and sometimes even significant variation within individuals or populations, leading to a broader circumscription of some species than that of previous workers. Incongruence between plastid and nuclear trees suggests the possibility of plastid introgression within section Hilsenbergia, and morphological data indicate potential hybridization, in areas of sympatry, among the three most widely-distributed species, D. baronii, D. cannabina, and D. hilsenbergii. Preliminary conservation assessments based on herbarium collections and field observations categorize D. wallichii as Critically Endangered and D. hafotsy as Endangered. This work summarizes the taxonomic history, morphology, nomenclature, and distribution of section Hilsenbergia and a dichotomous key to the species is provided.
Zuloagocardamum jujuyensis, a new genus and species of Brassicaceae from fujuy Province in Argentina, is described and illustrated, and its phylogenetic relationships to nearest relatives are demonstrated. It resembles some genera of the tribe Thelypodieae, such as Chilocardamum and Weberbauera, but differs mainly by having a well-developed woody caudex with reduced leafless stems, rosulate, awl-shaped or linear, sessile, parallel-veined basal leaves conspicuously ciliate with simple trichomes, racemes much shorter than the basal leaves, torulose fruits, and mucilaginous seeds. Phylogenetic analyses, based on DNA sequences of nuclear ITS and plastid ndhF and trnL-F regions, place Z. jujuyensis in the tribe Thelypodieae, where it is related to species of Weberbauera, Englerocharis, and Parodiodoxa. However, it is morphologically different from species of all four genera by the character combinations above.
Most recent treatments of Monotropsis recognize a single species of achlorophyllous, mycoheterotrophic herbs endemic to the southeastern U. S. A. (M. odorata), although four species have been described. This study reevaluates these proposed taxa using a comprehensive approach to species delimitation analyzing variation in morphology, phenology, geography, nuclear DNA, and chloroplast DNA. Principal components analysis of morphometric data reveals two clusters that are geographically distinct. These groups correspond to clades resolved with nuclear (ITS/26S and Xdh) and plastid (rpl32-trnL) DNA that are furthermore distinct phenologically. These data support recognition of M. reynoldsiae, a Florida endemic and M. odorata, an Appalachian endemic. Monotropsis lehmaniae, proposed based on an autumn flowering period and supposed floral differences, is shown to comprise individuals that are not yet in anthesis, and should not be recognized. Cryptophila pudica, named based on its disjunct geographical distribution and floral differences, is also shown to be indistinct. A taxonomic revision of the genus is presented.
Analyses of cpDNA using maximum parsimony, maximum likelihood and Bayesian inference reveal that Metastelma is polyphyletic in its present circumscription. Two species are transferred to Tassadia as T. richardiana and T. stricta. One species, M. eggersii, is retrieved as a member of subtribe Gonolobinae rather than Metastelmatinae, and several South American species fall within the “Stelmation” clade together with species of Ditassa. In Metastelma sensu stricto, the South American species are not monophyletic, falling into a M. tubatum Clade and a South American-Caribbean M. parviflorum Clade. Most Caribbean species, except for M. anegadense, M. decipiens, M. lineare, and M. parviflorum, form a clade in which the Cuban species are sister to the remaining taxa. The Central American mainland species show a clear geographical structure with a center of diversity in central Mexico, two radiations along the Occidental portion of Sierra Madre, and one radiation along the Oriental portion of Sierra Madre. Biogeographic analyses suggest that South America was the source for multiple invasions to the Caribbean. The genus possibly reached Central America via the Caribbean and the Yucatná peninsula, rather than the Panamanian land bridge, although further studies are needed to bolster this hypothesis. Several morphologically similar but geographically isolated taxa, interpreted previously as varieties and subspecies of the same species, are shown to be polyphyletic. As a result, Metastelma arenicola is raised to species rank, while M. trichophyllum, M. macropodum and M. chiapense are re-established as independent species.
Morphological data have produced conflicting results when analyzing evolutionary relationships within Gesneriaceae due to convergence of morphological characters. Columnea, the largest Neotropical genus in Gesneriaceae subfamily Gesnerioideae, has had a convoluted taxonomic history due to this convergence. Previously, the 200 species of Columnea were placed in 14 genera, with up to nine sections in the genus; most recently classifying the species in a single genus with five sections. Phylogenetic analyses presented here included 129 accessions representing 90 species for five chloroplast gene regions (trnQ-rps16 spacer, rpl32-trnLUAG spacer, rps16 intron, trnS-G spacer, and trnH-psbA spacer) and nuclear ribosomal ITS to build a well-supported topology that can test the previously proposed subgeneric classifications. Our goal was to test the monophyly of section Stygnanthe, one of the five sections that encompasses 18 species based on floral morphology. Fifteen species of Stygnanthe and four species that had not been classified in Stygnanthe but shared similar floral morphologies were included in the molecular phylogenetic analyses. Analyses indicate that the 19 species with a similar floral morphology belong in four separate clades including a newly proposed section, Columnea section Angustiflorae. Of the remaining three species with similar morphologies that were not sampled, only one is proposed to be a member of section Angustiflorae, none are members of clade Stygnanthe, but are likely to belong in other clades.
A new species, Plantago rahniana, is described and illustrated. This species has an extremely restricted distribution and is only known from two localities in Santa Catarina, in southern Brazil, which are high-elevation grasslands that are often covered in fog. Based on the IUCN criteria, this species is considered endangered. Material of this species was erroneously identified as Plantago commersoniana, which is also rare and potentially threatened. These two species mainly differ by the shape of their leaves and seeds, and by the pattern of the trichomes on their leaves, bracts, and anterior sepals. The first key to all of the species of Plantago from Brazil is also provided.
Species belonging to the genera Aloysia and Acantholippia are difficult to place within Lantaneae due to gene tree incongruence and limited sampling in previous studies. We use an expanded sample of both genera, and DNA sequence data from six loci, to reveal that Aloysia and Acantholippia species occur in five consistently inferred, well-supported clades. The precise relationships of these clades to one another are still enigmatic, due to gene tree incongruence. However, coalescent-based species tree inference supports the inclusion of most of Acantholippia in an expanded Aloysia sensu lato, with a 4-lobed calyx as its defining feature. Five new combinations are proposed to reflect this relationship: Aloysia deserticola, Aloysia riojana, Aloysia salsoloides, Aloysia tarapacana, and Aloysia trífida. Geographic range shifts from subtropical South America to North America have occurred at least twice in Aloysia. Shifts between determinate and indeterminate inflorescence arrangement have occurred at least twice independently. The elongate, lax inflorescence, which is characteristic of most of Aloysia, is hypothesized to be derived from a condensed inflorescence.
Vernonieae (Asteraceae) is a natural tribe defined mainly by discoid and homogamous heads, phyllaries imbricate in three to nine series, style branches acute with trichomes extending below the bifurcation, and cypselae 3–20-ribbed. It has been classified with morphological and molecular phylogenetic data into 21 subtribes, 127 genera and more than 1,000 species, distributed in the Americas, Asia, and Africa. The subtribe Lychnophorinae is almost completely restricted to the Central Brazilian Plateau with 11 genera, of which five are monotypic. It is usually defined by the lack of enlarged node at the base of the style and glands on the anther appendages, receptacle usually epaleaceous, and a pappus of capillary bristles or twisted straps. In the present paper we describe a new species, Anteremanthus piranii, and discuss its distribution and ecology. Morphological data were obtained from specimens collected during field trips (2011–2013) in the Serra Geral of Licínio de Almeida municipality, Bahia, Brazil. Anteremanthus piranii can be distinguished from the typical species A. hatschbachii mainly by the capitulescence terminal, 6–13 cm long (vs. axillar, 20–60 cm long), subinvolucral bracts linear at capitulum base (vs. subinvolucral bracts ovate), phyllaries pinkish-green (vs. greenish), florets 20–30, and corolla white, corolla lobes, anther, and style apex lavender (vs. florets 45–60, corolla green-cream). A distribution map, line drawing, and pictures of the new species are included.
The genera Lomatium and Cymopterus, along with many others, form a group that has been referred to previously as the perennial endemic Apiaceae subfamily Apioideae of western North America. This group of ecologically important and widespread species has been the target of numerous systematic studies, but the evolutionary relationships among these species remain elusive. Here we show that this confusion is due to high levels of morphological parallelism and homoplasy in the characters that have traditionally been used to define them, a result that is concordant with previous studies of the group. We explore patterns of evolution in traditionally important morphological characters using Bayesian stochastic character mapping on a phylogeny constructed from novel nrDNA and cpDNA sequence data for 96 specific and infraspecific taxa of the estimated 200 species in the group. We consider the implications of these results for taxonomic classification, the evolution of morphologies, and the utility of these morphologies to delimit small and large clades. Lomatium concinnum is newly combined as Cymopterus glomeratus var. concinnus and the new combination Cymopterus glomeratus var. greeleyorus is made.
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