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To test infrageneric classification and species delimitation within the pantropical moss genus Taxithelium (Pylaisiadelphaceae), we constructed a molecular phylogeny using three chloroplast loci (trnL, psbT and rps4), three mitochondrial loci (rps3, nad5 and nad4–5) and the nuclear marker ho1. Analyses of each locus separately and in various combinations, all support the monophyly of Taxithelium. Two major clades corresponding to taxonomically recognized subgenera were resolved within the genus. The first clade is composed of at least five smaller groups, four of which only include Southeast Asian plants and one is from the Americas and Africa; the latter is nested within the Southeast Asian groups. The second group appears to have a Southeast Asian origin with two dispersal events into America. Taxithelium is highly variable morphologically and includes plants with pluripapillose leaf cells as well as plants that lack papillae. Our data show that species T. nepalense, T. leptosigmatum, T. concavum, T. instratum, T. lindbergii and T. isocladum are each demonstrably monophyletic. On the other hand, T. planum and T. kerianum as circumscribed today are polyphyletic. The ho1 nuclear locus is used for the first time, with promising results for moss phylogenetic investigation.
KEYWORDS: Atlantic rainforest, Chromosome numbers, Endemism, lycophytes, South America, Floresta Atlântica, número cromossômico, endemismo, licófitas, América do Sul
Isoetes quiririensis is described, illustrated, and compared to similar species. This species can be distinguished from other species in Brazil by a set of characters that include microphylls that are semi-cylindrical in cross section, velum covering more than 3/4 of the sporangium, tuberculate megaspores varying from 477–670 µm in diameter, echinate microspores, and a chromosome number of 2n = 22. We also include a key for species from the Atlantic Rainforest that have tuberculate megaspores, distribution maps, and spore images for all species that are discussed. Isoetes quiriensis is known only from the type locality, where it occurs as an aquatic in small streams at high elevations. We suggest it should be classified as a data deficient species based on the IUCN criteria.
The morphologically variable tree fern Alsophila hornei of the Gymnosphaera clade is taxonomically and ecologically evaluated. While there is only one morphospecies present at the type locality on Fiji, three species can be characterized along an elevational transect on New Guinea. Alsophila hornei is also present at low elevations throughout New Guinea. Alsophila phlebodes is newly described and so far only found at high elevations on the Vogelkop Peninsula of New Guinea. Among its sympatric congeners, it stands out by having relatively pale brown petiole scales and well-developed, highly dissected remote basal pinnae on the petioles that form a wig-like structure around the trunk apex. Alsophila olivacea is reinstated and separated from A. hornei and A. phlebodes mainly by lacking adventitious pinnae and having an intermediate elevational distribution in the mountains of New Guinea. A key to the Australasian species of the Gymnophaera clade is provided in order to facilitate taxonomic studies of this biogeographically and phylogenetically interesting group.
The fern genus Osmolindsaea has recently been recognized as separate from Lindsaea on the basis of molecular data. Here we provide a taxonomic revision of Osmolindsaea based on morphological examination of herbarium specimens and molecular phylogenetic analysis of five plastid loci (trnL-tmF, trnH-psbA, rpoC1, rps4, and rps4-trnS). We recognize seven species in the genus, two of which are here described as new to science (O. latisquama and O. leptolepida) and three are transferred to this genus from Lindsaea (O. himalaica, O. minor, and O. plumula). A key to the species is provided, as well as descriptions, illustrations, synonymy, and a list of examined specimens for each species.
A vittarioid fern (Pteridaceae) distributed in Eastern Asia is described here as a new species, Haplopteris heterophylla, which has long been included in H. hainanensis. A description, illustration, distribution, ecology, and a key to congeneric relatives of the new species are provided. Obovoid-shaped paraphyses and frond dimorphism are the diagnostic characteristics of H. heterophylla. In a trnL-F matKndhF phylogeny of 14 Asian Haplopteris species, including H. hainanensis, both morphological observations and the cpDNA phylogeny support the systematic uniqueness of H. heterophylla.
Two new species of Diplazium from Brazil, D. adnatum and D. brachycarpum, are described and illustrated. A new name and status—Diplazium peruvianum—is proposed for Diplazium celtidifolium var. puberuletum. A list of characters differentiating these taxa and a key to the 2–3-pinnate-pinnatifid species of Diplazium from Brazil is provided.
Rumohra is represented by three species in Brazil. Two of them, R. glandulosissima and R. quadrangularis, are endemic and restricted to upper elevation openings in the Atlantic Rainforest from Rio de Janeiro and São Paulo States, and Rio de Janeiro and Rio Grande do Sul States, respectively. The third species, R. adiantiformis, is widespread. The new species here described, R. glandulosissima, is easily recognized by having densely glandular indument on all parts of the leaf. These glands are formed by two kinds of hairs. The first type are erect, capitate, ca. 0.1 mm long, hyaline, composed of three cells, with the basal and apical cells pale yellow, the central cell clear, and the apical cell slightly enlarged compared to the other cells. It does not produce any apparent exudate. The second type of glandular hair is sessile (dot-like), apparently formed by a single cell, ca. 0.1 mm wide, and less than 0.1 mm tall, and surrounded by a shiny reddish exudate. All three species are described, illustrated and distinguished by a key. These results are important for the conservation of rare plants in Brazil where there is a cottage industry of extracting leaves of Rumohra from wild populations for sale as floral greens.
A preliminary taxonomic revision is presented of the Neotropical genus Heteropsis, widely known as the source of a regionally important non-timber forest product; a tough resistant root fiber used in craftware and furniture making. Eighteen species and one variety are recognized, including two new species, Heteropsis reticulata and Heteropsis vasquezii; Heteropsis melinonii is considered to be of doubtful status. Descriptions, illustrations, geographical distribution and taxonomic commentary for each species, and an identification key are provided. Lectotypes are designated for Heteropsis oblongifolia, Heteropsis rigidifolia and Heteropsis salicifolia. Short reviews are provided of taxonomic history, phylogeny, vegetative and reproductive morphology, growth cycle, uses, and conservation status.
Vanilla labellopapillata, here described and illustrated, is a new species from the floodplain forest of Caxiuanã National Forest, situated in the northeastern Brazilian Amazon. The new species has consistent morphological features that support its inclusion in the Vanilla planifolia group. Vanilla labellopapillata is morphologically similar to V. insignis and V. odorata and diagnostic characters for separating it from the two species are given. A key to the Vanilla planifolia group is provided. Another species, V. cribbiana, recently described from Mexico and Central America is recorded for the first time in Brazil. Based on the examination of type material of Vanilla, we also propose the lectotipification of V. trigonocarpa and V. duckei.
Platanthera yadongensis, a new species of orchid from Yadong, Tibet, China, is described and illustrated. This new species is close to P. deflexilabella in sharing ciliate sepals, two long and outstretching stigmatic lobes, and naked and involute viscidia, but it differs from the latter in having bracts much longer than the ovaries in lower flowers, lips shorter than the sepals, spurs longer than the lips but much shorter than the ovaries and pedicels, and anther thecae slightly divergent towards the base. A key to all known species with two outstretching stigmatic lobes from China is provided.
About one hundred years ago, Dykes noticed an innovative feature in Iris, a raised central ridge along the midvein of the sepal, that he called a crest. Molecular phylogenetic and ancestral state reconstruction studies suggested that the sepal crest is a homoplastic character, even though the majority of the crested species form a monophyletic group. We investigated the putative multiple origins of sepal crests in Iris and relationships among crested species via comprehensive sampling of crested species in Iris using five plastid markers. We employed maximum parsimony, maximum likelihood, and Bayesian inference in reconstructing relationships. Our data analyses resolved a large core-crested clade along with four other independent lineages that also have crested species. Within the core-crested clade, four highly supported major clades were identified comprising species from subgen. Nepalensis and Scorpiris and some species from subgenus Limniris section Lophiris sensu Mathew. However, relationships among these four major clades were not fully resolved. Six species from section Lophiris represent four additional origins of the sepal crest.
Agavaceae subfamily Chlorogaloideae is composed of the North American genera Camassia, Chlorogalum, Hastingsia, and Schoenolirion, with many species occupying serpentine soils or other poor soils with unusual chemistries. The monophyly and intergeneric relationships of this group have not been rigorously assessed. We estimated the phylogeny of Chlorogaloideae using four chloroplast DNA regions: rpl16 intron and trnD-trnY-trnE-trnT, psbJ-petA, and trnS-trnfM spacers, with the goals of evaluating 1) the monophyly of Chlorogaloideae, 2) the monophyly of each genus and generic interrelationships, 3) the placement of Chlorogaloideae in Agavaceae, and 4) the history of adaptation onto serpentine soils. Maximum parsimony, maximum likelihood, and Bayesian analyses provided concordant estimates of the phylogeny supporting the monophyly of a clade consisting of Camassia, Chlorogalum, and Hastingsia, but suggest that Schoenolirion may be more closely related to Hesperoyucca and Hesperaloe. Each genus of Chlorogaloideae was found to be monophyletic except Chlorogalum, with C. parviflorum and C. purpureum forming a paraphyletic grade to other Chlorogalum, Camassia, and Hastingia. Ancestral character reconstructions employing parsimony, likelihood, and stochastic mapping suggest that serpentine tolerance was gained and lost multiple times in Chlorogaloideae with comparable rates of gains and losses with a small bias towards losses. We discuss the significance of the estimated phylogeny for the evolution of the distinctive bimodal karyotype of Agavaceae.
The circumscription of Brodiaea coronaria has remained in a state of flux over the past 200 yr. That species name historically was applied to a morphologically-diverse assemblage of populations ranging from southern California to coastal southern British Columbia. Recent floristic works recognized two subspecies, B. coronaria subsp. coronaria, which ranges from northern California to British Columbia, and B. coronaria subsp. rosea (currently = B. rosea), considered to be a serpentine endemic of northwestern California. A morphometric analysis was done to better refine the circumscription of both taxa. Ten floral characters and the length of the longest pedicel in the inflorescence of 10–50 were measured in plants from 63 populations of B. coronaria and five populations of B. rosea. Principal components analysis and cluster analysis were employed to determine whether discrete population groups could be recognized, and a discriminant analysis was done to further test these groups and to identify diagnostic characters. The results support a revised circumscription of B. coronaria, with many former populations of B. coronaria transferred to B. rosea. The analysis supports recognition of B. rosea at species rank, emended to include many non-serpentine populations. A new subspecies, B. rosea subsp. vallicola, is proposed for morphologically distinct populations that occur along the eastern edge of California's Central Valley. Despite this revision, B. coronaria remains a widely ranging and highly variable species, and further genetic and molecular data are needed to assess whether morphologically-cryptic polyphyletic lineages may be present.
A taxonomic revision of the grass genus Koeleria from the study of 3,255 specimens ranging from the western Mediterranean basin and Macaronesia is presented. We include descriptions and nomenclatural synonyms for all taxa inhabiting these regions. Detailed morphometric descriptions, illustrations, distribution maps, an identification key, and habitat data are given for each taxon. We recognize 10 species of Koeleria in this area, grouped into four sections. Four new combinations are provided for Koeleria, K. sect. Imbricatae (Domin) Quintanar & Castrov., K. sect. Reticulatae (Domin) Quintanar & Castrov., K. pyramidata subsp. arenaria (Dumort.) Quintanar & Castrov., K. pyramidata subsp. schroeteriana (Domin) Quintanar & Castrov., and one new name, K. rodriguez-graciae Quintanar & Castrov. Thirty-three names are lectotypified.
A new species of Guadua from the Atlantic forest in Argentina, Guadua variegata, is described and illustrated. Guadua variegata is compared to other taxa distributed in Argentina, Bolivia, Paraguay, Uruguay, and southern Brazil in a key and a table based on exomorphological characters. Additional micromorphological characters of foliage leaf blades and cauline epidermis of G. variegata are included. Lectotypes of G. paniculata and G. tessmannii are here designated.
Taxonomic features of species of Panicum section Parviglumia are reviewed and compared with those of other taxa in tribe Paniceae. The new genus Morronea, including six species ranging from Mexico to Argentina, is proposed on the basis of morphological and molecular characters. They are plants growing at forest edges, with membranous-ciliate ligules, lower glume reduced, 1/6 to 1/4 the spikelet length, lower palea and lower flower absent, and upper anthecium indurate with simple papillae and conspicuous macrohairs all over its surface. The new combinations: Morronea arundinariae, M. cayoensis, M. guatemalensis, M. incumbens, M. parviglumis, and M. trichidiachnis are proposed and the new genus is compared with other genera of the tribe Paniceae.
Chusquea purdieana is a previously poorly known species of woody bamboo (Poaceae: Bambusoideae: Bambuseae) from northern Colombia characterized by a scandent habit, a large number of leafy subsidiary branches per node, and narrow foliage leaf blades. A morphological analysis of similar collections from Venezuela and Colombia allowed us to distinguish two new species from the Venezuelan Andes, C. aurea and C. multiramea, based on a combination of vegetative and reproductive features. We here describe and illustrate the new species and we also provide a detailed redescription of C. purdieana, along with illustrations of its most diagnostic features. We report the occurrence of C. purdieana in Venezuela and document a broader range for the species in Colombia.
Dudleya (Crassulaceae) is a genus of succulent perennials endemic to western North America. Interspecific relationships within Dudleya have been difficult to discern due to a lack of synapomorphic characters for the genus or subgenera, highly variable morphologies within species, and extensive polyploidy. Here we present the first molecular phylogeny of diploid members of the genus using sequences from nrDNA and cpDNA. We cloned ITS alleles from six known polyploid individuals to determine their evolutionary origin. We have been able to resolve four clades within Dudleya, but many relationships are still uncertain. Our analyses support the monophyly of the genus and show that Dudleya is more closely related to North American Sedum species of the Sedoideae subfamily than to members of its currently circumscribed subfamily, the Echeverioideae. The current subgeneric distinctions within Dudleya are polyphyletic and should be revised in future taxonomic treatments. We tested the monophyly of several highly variable species and found that D. virens, D. cymosa, and D. abramsii are polyphyletic. The ITS alleles from polyploid taxa were not variable enough to resolve polyploid origins.
The endemic New Caledonian genus Podonephelium Baill. has representatives in all of the archipelago's main vegetation types (maquis, dry forest, and humid evergreen forest) and occurs on each of its principal substrates (calcareous, ultramafic and volcano-sedimentary). Recent phylogenetic studies confirm its monophyly and indicate that it forms a clade with two other endemic genera, Gongrodiscus and Storthocalyx. Podonephelium is characterized by juvenile leaves that form a whip-like structure and fruit that dehisce longitudinally into two halves that contain a round black seed nearly entirely covered by a red arillode. As part of a collaborative effort to clarify the taxonomy of New Caledonian Sapindaceae, a revision of Podonephelium is presented based on examination of all available collections coupled with extensive field studies. Nine species are recognized, four of which are newly described (P. cristagalli, P. davidsonii, P. pachycaule, and P. plicatum) and one infraspecific taxon is elevated to the rank of species (P. gongrocarpum). A key to species is provided, along with descriptions, distribution maps, line drawings (for the new species and new combination), and risk of extinction assessments using the International Union for Conservation of Nature IUCN red list criteria.
A new species, Hoya corymbosa, from Sabah (Borneo), is described and illustrated. It is a horizontally growing epiphytic shrub with diminutive flowers. Its minute staminal corona lobes lacking revolute outer margins and broad, flat caudicles of the pollinarium (resembling those of Dischidia) are quite peculiar within the genus, which is otherwise characterized by corona lobes with revolute margins and generally round to oval caudicles. Its position in Hoya has been confirmed by phylogenetic analysis based on nuclear ribosomal ITS region, the matK gene and psbA-trnH intergenic spacer.
A new species from St. Eustatius, Gonolobus aloiensis (Apocynaceae, Asclepiadoideae, Gonolobinae), is described and illustrated. This new species is endemic to St. Eustatius (northern Leeward Islands) and represents the first record of the genus for the island. It exhibits morphological similarity to continental G. albomarginatus, but can be distinguished in part by abaxial leaf surfaces glabrous, calyx and corolla lobes shorter and narrower, and abaxial corolla lobes completely lacking glandular indumentum and with eglandular indumentum restricted at most to only the top two-thirds of the lobes. An amplified key to species of Gonolobus s. s. in the West Indies is provided.
A new species from montane Colombia is described and illustrated: Gonolobus murphyae (Apocynaceae, Asclepiadoideae, Gonolobinae). The new species is morphologically similar to G. antioquensis (both exhibit caudate corolla lobes), but can be differentiated by a number of characters including vegetative pubescence, calyx pubescence, flower size, and shape of the laminar dorsal anther appendages.
The systematics and biogeography of the neotropical angiosperm genus Tachia Aubl. (Gentianaceae: Helieae) were investigated. The genus includes 13 neotropical species from Guyana in the east to western Peru in the west and from Costa Rica in the north to central Bolivia in the south. Tachia grandifolia var. orientalis is raised to the species rank as Tachia orientalis. Illustrations, a diagnostic key, phenology, distribution maps, and descriptions are provided for all species.
Cuscutatinctoria clade, the second largest infrageneric group of subgenus Grammica, includes 14 species that are centered in Mexico and adjacent regions, but also two species that are found in Australia, presumably as a result of long-distance dispersal. Flowers, pollen, and stigmas are among the largest in Cuscuta, with plants being xenogamous or facultatively xenogamous. In this clade, the convergent evolution of morphological traits, especially those associated with the calyx and gynoecium/ capsule, has obscured the identity of some species and has hindered previous efforts to determine their limits. Basic morphology, scanning electron microscopy and sequence data from the nuclear internal transcribed spacer (ITS) and the plastid trnL-F region were used to reconstruct the phylogeny, gain a better understanding of the evolutionary history, and determine species boundaries. Overall, species were grouped in five subclades, Based on their morphological and molecular similarity, C. tinctoria, C. aurea, and C. floribunda are best treated as a single species, with the latter two taxa being retained as varieties of the former. Our results also show that in their currently accepted delimitation, C. rugosiceps and C. tinctoria are polyphyletic, the former including one new species, C. volcanica , and the latter two new species, C. timida and C. tolteca. In addition, three other new species are described: C. iguanella and C. insolita, both with multicellular protuberances on the calyx and/ or corolla lobes, and C. montana, with broader than long calyx lobes. A taxonomic treatment that includes an identification key, descriptions, geographical distribution, ecological data, and illustrations for all taxa is provided.
Bupleurum dracaenoides, a new woody species endemic to southwestern China, is described and illustrated. This new species differs markedly in morphology from previously known congeneric species. It is characterized by its shrubby habit, with woody stems up to 1 m tall. The leaves are clustered at the top of the woody stems, the involucral bracts are ovate and unequal, the bracteoles are usually ovate or suborbicular and the pedicels of the mature fruits exceed the bracteoles. Molecular phylogenetic analyses indicated that Bupleurum dracaenoides belongs to a small subclade of subgenus Bupleurum and is sister to Bupleurum polyclonum. Relationships to other woody species of Bupleurum are discussed.
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