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The native goldback and silverback ferns of western North America, composing the genus Pentagramma, are phylogenetically isolated within the xeric-adapted cheilanthoid clade. Although species-poor compared to its sister group, Pentagramma encompasses a diverse array of morphotypes, cytotypes, and flavonoid chemotypes. Because the differences are generally cryptic, however, the various entities are usually recognized at an infraspecific level. In recent years, as many as five subspecies have been ascribed to P. triangularis, and only P. pallida has been considered sufficiently divergent to warrant recognition as a distinct species. In this study, we take a unified approach to taxonomic delimitation in Pentagramma. Combining spore studies with phylogenetic analyses of plastid and nuclear sequences, we identify six genetically and morphologically distinct diploid lineages, each of which is here treated as a species. A new species is described (P. glanduloviscida) and three new combinations are made (P. maxonii, P. rebmanii, and P. viscosa).
A vittarioid fern from the Solomon Islands, which has long been treated as Antrophyum semicostatum, is described here as a new species, Antrophyum solomonense. A description, illustration, and distribution of the new species are presented. Boot-shaped paraphysal apical cells, rounded stipe in cross section, and thin rhizome scale cell walls are the diagnostic characteristics of A. solomonense. A molecular phylogeny inferred from four chloroplast markers further supports its systematic uniqueness.
Moranopteris inaccessa, here described and illustrated, is a new species from southern Peru. Plants were collected in the Abra Málaga private conservation area from steep ledges accessible only with mountaineering equipment. Moranopteris inaccessa differs from its congeners by the combination of elongate membranaceous laminae that are 1-pinnate-pinnatifid. Found at 4,427 m, its distribution is over 500 m higher in elevation than any other species of Moranopteris. Phylogenetic analyses of atpß, rbcL, and trnL-trnF molecular sequences, using maximum parsimony, maximum likelihood, and Bayesian analysis support M. inaccessa as sister to M. longisetosa. The two are similar in their elongate and highly divided leaves compared to other species of Moranopteris. The discovery of this new species highlights the value of undertaking botanical surveys of undisturbed vegetation patches in the high Andes that potentially host unrecognized plant diversity.
Pinus douglasiana and P. maximinoi (Pinus subsection Ponderosae) are closely-related New World pines with vague taxonomic boundaries where their natural ranges overlap in western Mexico. They are distinguished from each other by the width of their leaves and thickness of their cone scale apophyses. They are also sometimes confused with two other close relatives, Pinus pseudostrobus and P. yecorensis. We integrated morphological, molecular, and ecological data to clarify the taxonomic limits among these four species. Following previous studies, we evaluated 16 quantitative leaf and seed cone characters. Pinus douglasiana, P. maximinoi, and P. pseudostrobus formed non-discrete groups in multivariate space. The absence of leaf hypodermal intrusions, a persistent peduncle, and the shape of the seed cone are useful for differentiating P. pseudostrobus and P. yecorensis from P. douglasiana or P. maximinoi, and the latter two can usually be distinguished by needle width or cone scale apophysis thickness. Most individuals identified as P. douglasiana, and P. maximinoi shared haplotypes for a plastid ycf1 fragment that is relatively variable for the genus, while P. yecorensis has a closely related, exclusive haplotype. A distinct haplogroup included all individuals of P. pseudostrobus and the remaining individuals of P. douglasiana and P. maximinoi. Leaf width and cone scale thickness of P. douglasiana and P. maximinoi are correlated with elevation. According to potential distribution models, P. yecorensis is distributed in drier areas than P. douglasiana or P. maximinoi, while P. pseudostrobus occurs in more temperate areas, commonly at higher elevations. Pinus douglasiana and P. maximinoi can be considered as incipient species undergoing divergent evolution characterized by incomplete morphological, molecular, and ecological divergence.
Aristolochia subgenus Siphisia comprises about 70 species, with a disjunct East- and South Asian to North- and Central American distribution. Here we describe and illustrate three new species of Aristolochia subgenus Siphisia from Vietnam: Aristolochia faviogonzalezii, A. tonkinensis, and A. tadungensis. The first two species are found in northern Vietnam while the latter occurs in southern Vietnam. Diagnostic characters distinguish the new species from their morphologically close relatives. Their distribution, ecology, phenology, ethnobotany, and conservation status are presented as well. In addition, three species are newly reported for the flora of Vietnam that were until now restricted to China, namely A. fangchi, A. hainanensis, and A. utriformis. An identification key to all Asian species of Aristolochia subgenus Siphisia is provided.
Three Neotropical orchid species, Mormolyca dodsonii, Mormolyca cruentata, and Mormolyca prunina are described on the basis of molecular and morphological characters. Phylogenetic relationships are estimated using sequence data from five plastid loci (rpoC1, matK gene and flanking trnK intron, atpB-rbcL intergenic spacer, and the 3′ portion of ycf1) plus the nuclear ribosomal internal and external transcribed spacers (ITS, ETS), and a portion of the low-copy nuclear gene phyC. Phylogenetic analyses confirm the monophyly of Mormolyca and clarify relationships within the genus: an early divergent paraphyletic grade of species diverging from the line leading to a major clade of two distinctive subclades. The three new species are contained within these subclades: M. dodsonii and M. cruentata clustering with the subclade characterize by flowers with a labellum pad of short glandular papillae; whereas M. prunina groups with the clade that includes all species with glossy labella.
Allium subgenus Melanocrommyum is taxonomically one of the most complicated groups within Allium and its sectional delimitation has been changed several times. In the present study, the relationships among 136 individuals representing 28 species of the sections Acanthoprason and Asteroprason were surveyed using ISSR (inter simple sequence repeats) markers. Nine ISSR primers generated 322 polymorphic bands (on average 36 bands per primer). In the resulting neighbor joining dendrogram, most conspecific individuals were grouped into clusters, which can be regarded as evidence of the monophyly of most species. Exceptions were the positions of six single samples of A. graveolens, A. zagricum, A. shelkovnikovii, A. subakaka, and A. materculae, which were not grouped with their remaining conspecific individuals. The clusters in dendrograms were significantly correlated with morphological similarities and biogeographical distributions. The lowest molecular similarity (62%) was observed between A. shelkovnikovii and A. ubipetrense and the most similar taxa (similarity value = 93%) were A. materculae and A. graveolens (sect. Acanthoprason). The results showed that ISSR analysis is suitable to detect the relationship of closely related species. The groups constructed agree only partly with those previously published based on molecular data, and represent alternative proposals to define taxonomic groups.
Genetic concepts within the Bromelioideae are highly problematic, in particular within the tank-forming “core bromelioid” clade. Previous molecular studies showed that the largest genus, Aechmea, and allied genera are polyphyletic and require revision. Here we focus on one group within the Aechmea alliance, the Portea/Gravisia group. To assess whether species of this group form a distinct lineage within the core bromelioid clade, and to clarify generic limits and interspecific relationships within the group, we generated and analyzed amplified fragment length polymorphisms (AFLP). In total, 69 species were sampled, including 26 species previously assigned to the Portea/Gravisia group. Neighbor joining, maximum likelihood and Bayesian analyses of the AFLP data consistently retrieved the Portea/Gravisia group as a monophyletic clade with several subclades, comprising species from the genera Aechmea, Canistrum and Portea. The phylogenetic distribution of polyporate pollen of the resulting trees indicates this character state arose once within the core bromelioids and thus can be regarded as a synapomorphy for the Portea/Gravisia clade. Further, our study shows that within the Portea/Gravisia group, subclades are characterized by petal color and geographic distribution. Thus, the present study is a further advance in the challenging task of elucidating phylogenetic relationships within the core bromelioids as the basis for a revised taxonomy of this ecologically important group.
One of the largest (ca. 450 accepted names) and most widely distributed genera of Bromeliaceae is Pitcairnia, with species occurring in Mexico, Central America, the West Indies, and throughout much of South America. The present study developed a phylogenetic hypothesis based on macro- and micromorphology, and tested the monophyly of the genus and subgenera, as well as the position of P. feliciana. The study included 67 species of Pitcairnia and 12 outgroup taxa selected to represent the morphological diversity and areas of endemism of Bromeliaceae. The results supported the monophyly of Pitcairnia, as indicated by two non-homoplasious and five homoplasious synapomorphies. Pitcairnia subgenus Pepinia and Pitcairnia subgenus Pitcairnia did not form a monophyletic group. Pitcairnia feliciana appeared to be related to the Atlantic Forest taxa. Phylogenetic relationships among the species recorded in the three Brazilian phytogeographic domains (Atlantic Forest, Cerrado, and Amazon) were obtained, and the relationships showed that the members of Pitcairnia occupied the three Brazilian domains through several routes.
Two new species and a new combination for Steyerbromelia (Bromeliaceae: Navioideae) are presented. These novelties represent the first records of Steyerbromelia for Colombia. The new species, Steyerbromelia naquenensis and Steyerbromelia nukakii, were placed in Steyerbromelia based on the combination of their 3- or 4-divided inflorescences, presence of smooth posterior sepals, and ovate, shortappendaged ovules. Both species resemble each other and S. thomasii, which also occurs in the basin of the Upper Rio Negro. Steyerbromelia garcia-barrigae is transferred from Navia, despite the lack of petal appendages, because its morphological characteristics fit better the limits of Steyerbromelia. An updated description of this species is presented. We now consider Steyerbromelia to be a genus of nine species. Additionally, notes on the geographic distribution, vernacular names, conservation status, and taxonomic comments of the three species are provided, as well as a key to all of the species. Finally, a discussion on the difficulties to place these three species within a genus is presented to highlight the problems of generic delimitation of the subfamily Navioideae.
Paspalum (Poaceae, Panicoideae) is a large genus comprising approximately 330 species, of which 210 are cited for Brazil. Paspalum minutispiculatum is similar to other related species of the informal “Humboldtiana” group, but this new species is annual and lacks rhizomes. This work establishes the distinction between this new species and related ones from this group, and contributes to a better characterization of the little known P. bicilium. Micromorphological characters of the upper anthecium and anatomical characters of the root, culm, and leaf of both species were analyzed. The new species appears to be endemic to the Federal District. It grows along trails in gallery forest, which is a rare habitat for this genus. The present work includes a key for species of the “Humboldtiana” group.
A new grass species from Brazil, Zizaniopsis longhi-wagnerae, is herein described, illustrated and compared with putative related species. The new species grows in swampy areas of montane grasslands of the southern plateau of Santa Catarina State, Brazil, and is thought to be endemic to this region. Comments about habitat, morphology, distribution, conservation status, and an updated key to the genus are presented.
Sabiaceae include over 120 species of mostly trees and shrubs found in moist forests of the Neotropics and Southeast Asia. The family includes three genera, Meliosma, Ophiocaryon, and Sabia, the first being the most species-rich and the only one found at the present time throughout the entire disjunct distribution range displayed by the family. Sabiaceae resolves among early-diverging eudicots in familylevel phylogenetic studies, however, to date, no such studies have focused on the family or any of its members. Nuclear and chloroplast molecular data were used to infer phylogenetic relationships within Sabiaceae and among species of Meliosma. Sabia occupies an early diverging position in Sabiaceae and Ophiocaryon is nested within Meliosma, although monophyly of the latter cannot be rejected. Results show that Meliosma alba, found in China and Mexico, is monophyletic. The infrageneric classification of Meliosma, largely based on endocarp morphological characters, is evaluated in light of these results. Findings suggest that the intricate floral morphology of Meliosma is derived within Sabiaceae and that the basic floral plan in the family is pentamerous. Results could support the transfer of Meliosma alba to a new genus if confirmed by future research.
A new species of Sauvagesia subsect. Vellozianae is described, Sauvagesia paganuccii, from material collected in campo rupestre vegetation of Bahia state, northeastern Brazil. The new species resembles S. insignis but differs mainly in its densely shrubby habit and smaller leaves and pedicel. Among the species of subsect. Vellozianae in Bahia, S. paganuccii is now the only one occurring in campos rupestres outside the Serra do Sincorá of Chapada Diamantina. An identification key to species of Sauvagesia subsect. Vellozianae from Bahia is also provided.
The genus Metrosideros (Myrtaceae) comprises 50–60 species found largely across the Pacific Islands. The relationships within this genus, including the circumscriptions of the subgenera Mearnsia and Metrosideros and their relationships with the other members of the tribe Metrosidereae (namely the New Caledonian endemic genus Carpolepis and the South American Tepualia), are poorly understood. Phylogenetic analyses were carried out using previously published ITS sequences, covering most species of the tribe, and new sequences of five single-copy nuclear genes with a reduced taxon sampling. The independent and combined analyses of the five nuclear genes using a range of approaches, including Bayesian single-gene, concatenated (MrBayes), concordance (BUCKy) and coalescent (*BEAST) analyses, yielded different topologies, indicating important conflicts among phylogenies based on individual genes. The deep relationships within the tribe Metrosidereae remain poorly resolved, but our results indicate that the species of Carpolepis and Tepualia are likely nested in the genus Metrosideros. A broad circumscription of the genus Metrosideros is therefore adopted, and the new combinations Metrosideros laurifolia var. demonstrans, Metrosideros tardiflora and Metrosideros vitiensis are here published.
A new species of Meriania (Melastomatacea; Merianieae) endemic to the Brazilian Atlantic Forest is described and illustrated. Meriania paratyensis is known only from a single locality in the Joatinga Ecological Reserve, municipality of Paraty, in Rio de Janeiro State. The diagnostic characteristics and comparisons with similar species, geographic distribution, and a conservation assessment using IUCN guidelines are presented for the new species, as well as a key to identify all Meriania species known from the Atlantic Forest.
Davidia involucrata Baill. (Davidiaceae), commonly known as “Chinese dove tree”, is a rare and endangered species endemic to China. Little is known about the genetic structure and population history of D. involucrata. Both cpDNA and nSSR markers showed high genetic diversity among D. involucrata populations (HS = 0.717; hT = 0.893; pT = 3.14 × 10-3). The genetic divergence of all D. involucrata populations was significantly higher in the cpDNA (FST = 0.765) than in the nuclear DNA (FST = 0.239), possibly reflecting a more important role of pollen dispersal than the seed in the formation of the genetic structure of Davidia. Phylogeographic analyses suggested that the populations of D. involucrata were divided into the eastern and western lineages with the Sichuan Basin as the boundary, separating the Sino-Himalayan and Sino-Japanese Forest subkingdoms. Divergence of the two lineages was estimated to have occurred around 4.81 Mya during the Late Miocene/Early Pliocene. Mismatch distribution and neutrality tests didn't show significant population growth and range expansion in either lineage, suggesting the in situ survival of multiple populations during the Quaternary glaciation without range expansion. There are two potential refugia of Davidia involucrata in Southwest and Central China and natural reserves need to be established to preserve the genetic diversity of the species in the refugia.
Sabatia (Gentianaceae) contains ca. 20 species, distributed mainly on the U. S. A. Gulf and Atlantic Coastal Plains. Our aims were to determine 1) phylogenetic relationships among Sabatia species, 2) the time and place of Sabatia's origin and main areas of diversification, 3) relationships among sympatric species, and 4) how morphological and karyological characters evolved. We sequenced five noncoding cpDNA regions and nrITS for 30 accessions of Sabatia, Gyrandra, and Eustoma. Parsimony, likelihood, and Bayesian phylogenetic analyses were performed. Bayesian dating was done on a reduced-taxon, combined molecular dataset. The maximum clade credibility chronogram was used for ancestral area reconstruction and character optimization. Correlations between distributional, environmental and phylogenetic matrices were tested with spatial analyses. Phylogenetic analyses reveal that a SabatiaGyrandra clade diverged in the late Middle Miocene, with Sabatia subsequently splitting into western and eastern Gulf Coast clades during the early Late Miocene. Further diversification took place in the Late Miocene-Pliocene, with more recent range expansion. Pliocene glacial/interglacial periods could have triggered range contraction/ expansion, associated with chromosomal changes. Closely related species of Sabatia tend to share both distributions and habitat types. Character optimization showed potential synapomorphies for a polymerous clade and a white-flowered clade.
In the California Floristic Province (CA-FP) and nearby deserts, 19 species of Salvia (Lamiaceae, Mentheae) form a small radiation but an important component of the chaparral and desert communities. Traditionally, two groups within these Californian Salvia have been recognized (usually treated as sections), but relationships within each, to each other, and to other Salvia are unclear. Phylogenetic relationships of all species, with multiple accessions for most, were obtained using chloroplast DNA (cpDNA) and nuclear ribosomal DNA (nrDNA) markers. Ancestral character state reconstruction of both vegetative and floral features was done on the resulting nrDNA tree. Biogeographical analysis of the groups within the CA-FP and adjacent floristic provinces was done in BioGeoBEARS and species diversification assessed with BAMM. Significant conclusions drawn from the study include: 1) California Salvia should be classified into two monophyletic sections, Audibertia (15 species) and Echinosphace, (four species) in the new subgenus Audibertia; 2) subg. Audibertia and the Neotropical subg. Calosphace are sister clades, most closely related to Asian groups, and are likely Asian in origin; 3) nrDNA provides a fairly resolved tree for subg. Audibertia with all species monophyletic; 4) cpDNA and nrDNA trees are strongly incongruent and provide evidence that hybridization and chloroplast capture have played an important role in the evolution of subg. Audibertia; 5) ancestral character reconstruction of states in habit, possession of spines, calyx lobing, and staminal features highlights a complex (sometimes convergent) evolutionary history of this iconic CA-FP lineage; 6) subg. Audibertia arose in desert areas and more recently diversified into the southwestern California region and adjacent regions with the formation of the Mediterranean-like climate; and 7) this diversification exhibits a slight decrease in speciation and an increase in extinction rates over the group's 11 million year history.
Diagnostic seed surface morphologies have been identified in some members of the family Plantaginaceae; however, limited diagnostic seed morphologies have been identified in the subtribe Gratiolinae of tribe Gratioleae. Tribe Gratioleae consists of 25 genera and about 182 species distributed worldwide. Ten genera are classified under subtribe Gratiolinae. The current study employed scanning electron microscopy (SEM) to examine detailed surface sculpturing in the subtribe Gratiolinae to identify diagnostic seed characters of taxonomic significance. Seeds of 35 species representing seven genera were carefully sampled from herbarium specimens, mounted on aluminum stubs, observed under the SEM, and analyzed. Seeds were typically less than 1.0 mm in size. Eleven reticulate seed types were identified with significant variation. Variation resulted from at least five characters: overall seed pattern, seed length, periclinal wall ornamentation, anticlinal wall relative height, and ornamentation. Most seed types were characteristic of genera. Seed types of Bacopa did not correspond to sectional treatment of the genus and seed type observed for the monotypic segregate genus Benjaminia was shared with some members of Bacopa. Seed types identified for the segregate genus Mecardonia were not observed in Bacopa, whereas similar seed types were observed between some members of Mecardonia and the monotypic genus Amphianthus. The diversity in seed surface morphology suggests extensive but taxonomically significant seed variation in the subtribe Gratiolinae.
A new taxonomic circumscription of Solanum section Lathyrocarpum is proposed to include 11 species from North and South America. The section is placed in the cosmopolitan S. subgenus Leptostemonum, which contains 350–450 species. As defined here, S. sect. Lathyrocarpum includes all the taxa resolved in the Carolinense clade in previous molecular phylogenetic studies as well as one species not sampled in those analyses. The combination of morphological characters that define the group includes: herbaceous perennial habit with extensive root systems, often with endogenous buds that can sprout, a sparse to dense indumentum of sessile to stalked stellate hairs and tapered prickles, 2- to 3-foliate to plurifoliate sympodial units, extra-axillary inflorescences with 1–20 flowers, a weakly andromonoecious reproductive system, corollas ranging in color from white to lavender, and fruits maturing to yellow or yellowish-orange. Most species in the section have a weedy habit and grow in highly disturbed areas, but four are narrowly distributed. One species, S. carolinense, is a noxious invasive weed that has been inadvertently introduced to many areas around the world. There are a variety of ploidy levels represented in the group (n = 12, 18, 24, and 36). In this work, we present a new circumscription of the section and provide descriptions, distribution maps, preliminary IUCN conservation assessments, and a key to identify the 11 included species. We also designate lectotypes for nine names and select a neotype for S. dimidiatum.
P. López-Sepúlveda, K. Takayama, J. Greimler, D. J. Crawford, P. Peñailillo, M. Baeza, E. Ruiz, G. Kohl, K. Tremetsberger, A. Gatica, L. Letelier, P. Novoa, J. Novak, T. F. Stuessy
Oceanic islands provide many opportunities for examining modes of speciation in endemic plants, especially using modern molecular methods. Most speciation has been by either cladogenesis, usually resulting in radiated complexes, or anagenesis, yielding single transformed species. Previous genetic studies with available molecular markers, representing a limited sampling of the genome, have suggested that during cladogenesis, species accrue small genetic differences but differ dramatically in morphology. An appropriate archipelago in which to evaluate the genetic consequences of cladogenesis is the Juan Fernández Islands, a national park of Chile. This present study focuses on AFLP and SSR genetic differences among and within populations of six endemic species of Erigeron (Asteraceae), restricted principally to the younger island, Alejandro Selkirk (one to two million years old). Results show that three different genetic lineages exist among these species: (1) E. rupicola and E. stuessyi; (2) E. fernandezianus; and (3) the E. ingae complex (including E. ingae, E. luteoviridis, and E. turricola). The three genetic lines are distinct from each other, and each harbors considerable genetic variation. The three species of the E. ingae complex appear to be segregating genetically but are not yet ecologically divergent. The amount of genetic differentiation among species of Erigeron is less than that already documented among species of Robinsonia (Asteraceae) on the older island (four million years old). Erigeron fernandezianus, occurring on both islands, appears to have originated on Alejandro Selkirk Island and subsequently dispersed and established also on Robinson Crusoe Island.
The taxonomic complexity of the Senecio doria group (Compositae, Senecioneae) is reflected in the heterogeneous treatments that have been proposed and the poor knowledge of some species. This species group consists of perennial herbs distributed in Europe, western and central Asia, and northwestern Africa. The first worldwide revision of this group recognizing seven species is presented here (i.e. S. altissimus, S. bithynicus, S. doria, S. fontanicola, S. legionensis, S. morisii, and S. umbrosus). In this new taxonomic treatment S. legionensis and S. fontanicola are recognized at the specific rank. On the other hand, S. macrophyllus from eastern Europe/western Asia has been synonymized to S. doria. Senecio bithynicus and S. morisii are described as new species. Eleven names are lectotypified. Descriptions, distribution maps, and an identification key are provided for all the species included.
Two new species of Burmeistera (Campanulaceae: Lobelioideae) from the Cordillera de Talamanca are described, illustrated, and discussed with reference to similar species. One species, B. serratifolia, is endemic to Panama, while the second, B. monroi, is known from both Panama and Costa Rica. Additionally, these species are included in a dichotomous key to all Central American species of Burmeistera.
Valeriana serratifolia, a new species from the Andean region of northwestern Argentina, is described and illustrated. It can be distinguished from its closest morphological taxon, V. macrorhiza, by its entire, spathulate leaves with serrulate margin, and falcate fruits, 2–2.5 mm long, crowned by the reduced annular calyx shifted toward the ventral face. A phylogenetic analysis based on DNA sequences of plastid markers trnL-F and trnK intron supported V. macrorhiza as its closest related species. We also provide a key to differentiate V. serratifolia from the epappose species of Valeriana from Argentina.
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