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Microsatellite markers were used to test whether two recently described species of Sphagnum (Bryophyta), S. atlanticum R.E. Andrus and S. bergianum R.E. Andrus, represent distinct gene pools. The first species is considered endemic to eastern North America while the second species has been reported from Alaska and Newfoundland. The results indicate that S. atlanticum does not differ genetically from the closely related species, S. torreyanum, also restricted to eastern North America. In fact, some samples that are identical across all 15 micro-satellite loci have been distinguished morphologically as these two species. Plants of S. bergianum from Alaska are closely related genetically to Alaskan plants of the similar species, S. subfulvum, whereas Newfoundland plants of S. bergianum are more closely related to Newfoundland plants of S. subfulvum. Alaskan versus Newfoundland plants of S. subfulvum s.l. (including S. bergianum) are differentiated at microsatellite loci. Another closely related species, S. subnitens, is distinct from S. subfulvum and S. bergianum. Sphagnum atlanticum is synonymized under S. torreyanum and S. bergianum is synonymized under S. subfulvum.
A new fern species, Polystichum weimingii (sect. Metapolystichum, Dryopteridaceae), is described from southern Yunnan, China. The new species is morphologically most similar to P. discretum. Compared to P. discretum, P. weimingii has a smaller habit, lanate (linear and curly), brown rachis scales, and pinnules without obvious aristate spines on the margin. In contrast, P. discretum is larger, has linear, dark brown rachis scales, and has pinnules with obvious aristate spines on the margin. The new species also differs from P. discretum in spore morphology. Features of the new species in anatomy of lamina epidermis, spore morphology, geographical distribution, and ecology are given. The relationships with three other closely related species in Polystichum ser. Linearia are discussed.
Guided by systematic studies on Melpomene, we tested the monophyly of the genus by analyzing sequences of cpDNA (atpβ, rbcL) and morphological characters for 24 of the 29 recognized species. The atpβ data did not yield any resolution of the investigated group. The other data sets each supported the monophyly of the genus, as did all data combined, and indicate it to be sister to the Lellingeria apiculata group as proposed in previous studies. While the morphologically distinct M. leptostoma and M. gracilis are basal lineages with long branch lengths, most of the relationships within the core genus are weakly supported and have very short branch lengths. This indicates a recent radiation of the genus in the northern Andes, its center of diversity, apparently under the influence of the fast uplift of the mountain range and of glacial fluctuations of the vegetation zones, with subsequent dispersals to the Guyana Highlands, the Brazilian shield, and Africa.
A new species, Smilax pilosa, is described from the Serra do Brigadeiro State Park, in Minas Gerais State, Brazil. Smilax pilosa is the only Brazilian species of this genus having a dense indumentum covering the entire abaxial surface of the leaf. The new species is similar to Smilax muscosa Toledo but differs in that the latter has only a sparse indumentum restricted to the veins of the abaxial leaf face. The present study examines the affinity between the two species, reports the newly extended geographic distribution of Smilax muscosa into Espirito Santo State, and presents a key to the 22 species of Smilax that occur in Minas Gerais.
Paspalum, an American genus of the x = 10 Paniceae clade, includes about 330 species, four subgenera, and 27 informal groups. Within the genus, the Quadrifaria and Virgata groups are well-represented in South and Central America. Interspecific variability make the delimitation of these groups difficult; hence several species have been included or excluded from Quadrifaria or Virgata depending on the taxonomic treatment. In previous analyses, the Quadrifaria and Virgata groups of Paspalum were polyphyletic. Here we present a new appraisal of the classification of both groups based on the phylogenetic analyses of DNA data from the chloroplast: the rpl16 intron, and the region comprising the trnL and trnF genes. A monophyletic Virgata clade is recovered, consisting of nearly all the species listed in an unpublished manuscript. The Quadrifaria group is restricted to P. quadrifarium and P. quarinii. Other clades grouped species traditionally treated within Virgata or Quadrifaria, although their phylogenetic placement needs to be reevaluated. Within most of the clades, diploid and polyploid species seemingly derive from a common ancestor denoting an autopolyploid origin. Alloploidy is also possible although reticulate evolution needs to be explored. The wide distribution of the I genome suggests that this is a plesiomorphic state in Paspalum.
Menispermaceae, a dioecious, largely pantropical family of vines and lianas, consists of approximately 70 genera. In this paper, we present phylogenies (derived using maximum parsimony, maximum likelihood, and Bayesian inference) based on chloroplast atpB and rbcL sequence data for 42 Menispermaceae genera (60 species) and 15 outgroup genera representing all families in the Ranunculales. Results of this work confirm that a clade consisting of Berberidaceae and Ranunculaceae is sister to the Menispermaceae and that Menispermaceae is highly supported as monophyletic. Within Menispermaceae, Menispermum and Sinomenium, two genera with distributions in the Northern Hemisphere, are weakly to moderately supported as sister to all remaining Menispermaceae. Many of the tribes as described by previous workers are not monophyletic. To expand both the data and sampling, we combined our atpB and rbcL data with previously published ndhF data for two additional analyses: 1) for matching species and 2) for all genera available, including those with either atpB/rbcL or ndhF missing (22% missing data). All trees produced from our three analyses were highly congruent, especially where branch support was high. Four major clades were recovered from all analyses and are informally named. Superimposing our geographical distributions on our atpB/rbcL phylogeny indicates that many well supported clades have distributions on multiple continents, probably due to both vicariance and dispersal. Several previous hypotheses regarding trends in character evolution are tested; our phylogenies suggest that the following are all derived character states within the family: arborescent habit; acrodromous, peltate, compound, or lobed leaves; absence of condyle and endosperm; smooth endosperm; fused stamens; and tricolpate pollen.
The Amaranthaceae-Chenopodiaceae alliance has been the focus of several phylogenetic studies, but major questions concerning the internal relationships of Amaranthaceae still remain unanswered. This study aims to test the monophyly of the subfamily Gomphrenoideae and to examine the generic relationships within this group. Parsimony and Bayesian analyses of trnL-F and rpl16 show that the subfamily Gomphrenoideae is monophyletic. The clade of Gomphrenoideae represents a large and diverse radiation of the Amaranthaceae in the New World. Unilocular anthers are a morphological synapomorphy for Gomphrenoideae that was derived from bilocular anthers. Three major clades are resolved: the Gomphrenoids, the Alternantheroids, and the Iresinoids, which are largely supported by pollen morphology. The Iresinoid clade is sister to Alternantheroids plus Gomphrenoids, rendering metareticulate pollen as the synapomorphy for the latter two clades. Tribes and subtribes delimited by androecium and inflorescence characters are poly- or paraphyletic. Several genera are monophyletic including the large genus Alternanthera, whereas Gomphrena is polyphyletic. Irenella and Woehleria are resolved within Iresine whereas Blutaparon and Lithophila fall within the polyphyletic Gomphrena. The trnL-F and rpl16 sequence data are the most variable chloroplast regions examined to date for the family and are highly effective in resolving relationships in Amaranthaceae.
A comprehensive taxonomic revision of the genus Pachycereus (Cactaceae, Cactoideae, Pachycereeae) is presented. Pachycereus is characterized by one synapomorphic character, the presence of a narrow interareolar groove, and by the combination of flowering region differentiated with flexible spines, flower with a narrow region between the pericarpel and the receptacular tube and the occurrence of trichomes and short spines in the pericarpel. Five species are recognized in the present treatment. One species (P. pecten-aboriginum) is widely distributed from Chihuahua in northern Mexico to Chiapas, three species (P. grandis, P. pringlei, and P. weberi) are restricted to defined floristic provinces, and one species (P. tepamo) is endemic to Depresion del Balsas. A key and descriptions of the species, based on herbarium specimens, field observations and intensive field collection are presented. The taxonomic history, synonymies, distribution, uses, and representative specimens are included. Three new combinations, Pterocereus gaumeri subsp. foetidus, Lemaireocereus lepidanthus, and Lophocereus marginatus, are proposed.
Twelve Caribbean species of Vachellia (formerly Acacia subgenus Acacia) are formally monographed with taxonomic, habitat, geographic distribution descriptions, and lists of representative specimens examined. The species treated here are all part of the informal Vachellia acuifera group, which is characterized by generally enlarged stipular spines borne in clusters on the stems, 1–3 pairs of pinnae per leaf, and 20–50 stamens. Ordination analyses of vegetative characters show that these species form discrete groups in two- and three-dimensional plots. Vachellia baessleria Clarke, Seigler, & Ebinger, endemic to Guantánamo and Las Tunas, Cuba, is newly described. Vachellia acuifera is wide ranging, occurring from the Bahamas to Cuba. The other species are highly restricted in their distribution and ecological setting. Four of the species are only known from Hispaniola (V. barahonensis, V. caurina, V. cucuyo, and V. oviedoensis) and six only from Cuba (V. baessleria, V. belairioides, V. bucheri, V. daemon, V. roigii, and V. zapatensis). Another species, V. anagadensis, is only found on the island of Anegada in the British Virgin Islands.
The order Fabales, including Leguminosae, Polygalaceae, Quillajaceae and Surianaceae, represents a novel hypothesis emerging from angiosperm molecular phylogenies. Despite good support for the order, molecular studies to date have suggested contradictory poorly supported interfamilial relationships. Our reappraisal of relationships within Fabales addresses past taxon sampling deficiencies, and employs parsimony and Bayesian approaches using sequences from the plastid regions rbcL (166 spp.) and matK (78 spp.). Five alternative hypotheses for interfamilial relationships within Fabales were recovered. The Shimodaira-Hasegawa test found the likelihood of a resolved topology significantly higher than the one calculated for a polytomy but did not favour any of the alternative hypotheses of relationship within Fabales. In the light of the morphological evidence available and the comparative behavior of rbcL and matK, the topology recovering Polygalaceae as sister to the rest of the order Fabales with Leguminosae more closely related to Quillajaceae Surianaceae, is considered the most likely hypothesis of interfamilial relationships of the order. Dating of selected crown clades in the Fabales phylogeny using penalized likelihood suggests rapid radiation of the Leguminosae, Polygalaceae, and (Quillajaceae Surianaceae) crown clades.
Generic level relationships in phaseoloid legumes have received much attention using chloroplast DNA markers. However, despite this attention not all relationships are yet well-resolved. This study includes trnL-F sequences from across a wide sample of phaseoloid legumes as well as seven additional chloroplast DNA loci (rbcL, atpB, trnK/matK, rpl2, clpP, rps16, and ycf4) analyzed separately and in combination. Together, these data provide support for many relationships generally consistent with, but only weakly supported, in earlier studies. Some major discordant phylogenetic results were found in our separate analyses; for example, ycf4 sequences group Glycine and Teramnus with strong support; however, the combined analysis of the remaining seven loci found incongruent groupings (Glycine and Psoraleeae genera; Teramnus and Amphicarpaea) also with strong support. Network analysis of ycf4 revealed that the conflicting signal (relative to the other seven loci) came from first and second codon positions. These positions also showed significant rate acceleration, together indicating that selection driving convergent molecular evolution is the likely cause of the signal in ycf4, rather than shared history. The major clades within the phaseoloid legumes supported by our analysis are discussed.
The greater Caribbean region has played an important role in the early diversification of Croton L. (Euphorbiaceae s.s.). Jamaica is also important to Croton taxonomy because several of the earliest described species were based on material from the island. The Jamaican species of Croton are found in five distinct clades indicating that there were at least five separate over-water dispersal events of the genus to the island. Croton jamaicensis (section Eluteria Griseb.), a new species endemic to limestone hills along the southern coast of Jamaica, is described and illustrated. The species is phylogenetically most closely related to C. laurinas Sw. and C. grisebachianus Müll. Arg., both also endemic to Jamaica. Several lectotypifications and novel synonymies are required to clarify the taxonomy of the species of Croton that have been described from Jamaica, some of which are widespread in the region.
A phylogenetic analysis was conducted on the tribe Spiraeanthemeae (Cunoniaceae) to clarify relationships of Acsmithia and Spiraeanthemum. Three molecular markers, one plastid region (trnL intron and trnL-trnF intergenic spacer) and two nuclear single copy genes (ncpGS and PHYC), were sequenced for this purpose. The independent analysis of the three markers and a combined analysis all showed that Acsmithia is paraphyletic since Spiraeanthemum is nested within it. A morphological survey of all species in the tribe confirmed the existence of two groups within Acsmithia. One comprises the species from Australia, New Guinea, and A. densiflora from New Caledonia and is characterised by multiple ovules per carpel. The other group contains all the remaining New Caledonian species plus A. vitiensis from Fiji and is characterized by a single ovule per carpel. The study shows that characters previously used to distinguish Acsmithia and Spiraenthemum, phyllotaxy and sexual system, are homoplasious as in several other groups of Cunoniaceae. A broad circumscription of the genus Spiraeanthemum is adopted here that includes the species formerly placed in Acsmithia. Two new combinations are proposed: Spiraeanthemum collinum and Spiraeanthemum meridionale. Spiraeanthemum austrocaledonicum is considered a synonym of Spiraeanthemum densiflorum.
The genus Polemonium is composed of animal-pollinated herbs found largely in North America. Wide phenotypic variation suggests rapid evolution of floral and vegetative morphology. However, phylogenetic relationships within Polemonium have received little attention. We present a phylogeny of Polemonium based on 1676 Amplified Fragment Length Polymorphisms (AFLPs). We obtained similar topologies through minimum evolution analysis using Nei-Li genetic distances and parsimony analysis. Our analyses supported the mono-phyly of most species, and indicated that the single annual species, P. micranthum, is sister to all perennial species. Relationships were well resolved among species separated by genetic distances<0.16. Three well-supported clades each included taxa from one of three species complexes: the P. caeruleum complex, the P. pulcherrimum complex, or the P. viscosum complex. Relationships within clades were generally consistent with earlier systematic work. Exceptions were the inclusion of eastern North American P. reptans within the P. caeruleum clade, inclusion of P. boreale within the P. pulchernmum clade, and a close relationship between tetraploid P. califomicum and diploid P. pulchernmum subsp. delicatum. Our results indicate that floral traits are quite divergent among close relatives within the genus, and provide a phylogenetic framework for research on adaptation and speciation in Polemonium.
A phylogenetic study and taxonomic revision of the four currently accepted species of Bartonia (Gentianaceae, sub tribe Swertiinae) were conducted in order to test species boundaries and interspecific relationships. Species boundaries were examined based on measurements of key quantitative and qualitative morphological characters as given in the original descriptions. Phylogenetic analyses were performed using molecular data from the nuclear internal transcribed spacer region and chloroplast DNA (trnL intron through the trnL-F spacer), separately and combined using parsimony and Bayesian methodologies, incorporating outgroups from subtribes Swertiinae and Gentianinae. The morphological study revealed that characters of one species, B. texana, represent a subset of the morphological variation found within B. paniculata, but that B. paniculata, B. verna, and B. virginica could all be separated from one another. The molecular phylogenetic analyses all found B. texana to nest in a clade with the two recognized subspecies of B. paniculata (subsp. paniculata and subsp. iodandra), making the latter paraphyletic. Bartonia texana is here reduced to subspecific rank, as Bartonia paniculata subsp. texana. Also, the phylogenetic analyses showed strong support for a sister group relationship between B. verna and B. virginica, as opposed to between B. paniculata and B. virginica as has been previously suggested.
The genus Leucas R.Br. includes 100 recognized species ranging in distribution from Africa through the Indian subcontinent to Queensland, Australia. Morphological cladistic studies have suggested that several other genera of Lamioideae - Acrotome, Isoleucas, Leonotis, and Otostegia (pro parte) - may be closely related to Leucas, or even nested within it. Here we use phylogenetic analysis of three plastid DNA loci, the trnL-F region, trnS-G spacer, and the rps16 intron and reach similar conclusions. Many of the morphological features used to maintain Leucas since 1810 are shared ancestral (symplesiomorphic) characters. The other genera (or subgeneric group in the case of Otostegia) are therefore defined by apomorphic states. This is particularly apparent in the case of Leonotis, with its highly specialized bird-pollinated flowers, derived within a paraphyletic and insect-pollinated Leucas. Some geographically isolated groups are monophyletic, including all Asian species of Leucas, a Socotran clade, and a large African lineage.
Recent molecular systematic research has indicated the need for a revised circumscription of generic boundaries in subtribe Castillejinae (tribe Pedicularideae, Orobanchaceae). Based on a well-resolved and well-supported phylogenetic hypothesis, we present a formal reclassification of the major lineages comprising the Castillejinae. Prior to this treatment, subtribe Castillejinae included Castilleja (ca. 190 spp.), Cordylanthus (18 spp.), Orthocarpus (9 spp.), Triphysaria (5 spp.), and the monotypic genera Clevelandia and Ophiocephalus. In the classification presented here, Orthocarpus and Triphysaria retain their current circumscriptions, Castilleja is expanded to include Clevelandia and Ophiocephalus, and Cordylanthus is split into three genera; a key to the genera as they are recognized here is provided. Two new combinations, Castilleja beldingii and Castilleja ophiocephala, are proposed within the expanded Castilleja. The concept of Coriylanthus is restricted to the 13 species formerly recognized as subg. Coriylanthus, while subg. Dicranostegia and subg. Hemistegia are elevated to genus level (Dicranostegia and Chloropyron, respectively). We resurrect the generic name Chloropyron for the halophytes previously recognized as subg. Hemistegia. Five new combinations are proposed for Chloropyron (Chloropyron maritimum subsp. canescens, Chloropyron maritimum subsp. palustre, Chloropyron molle subsp. hispidum, Chloropyron palmatum, and Chloropyron tecopense). In addition to the formal classification, we provide phylogenetic clade definitions for Castillejinae, each of the genera, and two additional clades that are not assigned formal ranks. Morphological characteristics used to recognize traditional groups are evaluated, and synapomorphies are discussed. Finally, the current infrageneric classifications for Castilleja and Coriylanthus are evaluated in light of the recent molecular phylogenetic analyses.
Nierembergia (Solanaceae) contains 21 species found primarily in South America, with one species occurring disjunctly in Mexico. Among other features, a pollination system not found elsewhere in the Solanaceae typifies this genus. In most species, nectaries are absent, but oil-producing elaiophores are present on the corolla limb and these attract oil-collecting bees. Molecular phylogenetic hypotheses based on nuclear ITS and chloroplast rpl16 intron data support the monophyly of Nierembergia, which is sister to a clade composed of Bouchetia plus Hunzikeria. The data reconstruct two clades within the genus: one composed primarily of herbaceous species with broad stigmas and the second containing woody species with crescent-shaped stigmas. Morphological homoplasy is frequent among the species of the herbaceous clade, while the woody clade is morphologically cohesive. Chromosome counts are confirmed and karyotype features are given for 13 Nierembergia species (including six varieties) and for Bouchetia anomala, Leptoglossis linifolia, and Petunia axillaris. The two clades are also supported by karyotypic features: one has asymmetrical karyotypes, small chromosomes, and two chromosome pairs with nucleolar organizing regions (NOR) and the other has symmetrical karyotypes, large chromosomes, and only one NOR.
Solanum section Petota is taxonomically difficult, partly because of interspecific hybridization at both the diploid and polyploid levels. There is much disagreement regarding species boundaries and affiliation of species to series. Elucidating the phylogenetic relationships within the polyploids is crucial for an effective taxonomic treatment of the section and for the utilization of wild potato germplasm in breeding programs. We here infer relationships among the potato diploids and polyploids using nitrate reductase (NIA) sequence data in comparison to prior plastid phylogenies and: 1) examine genome types within section Petota, 2) show species in the polyploid series Conicibaccata, Longipedicellata, and in the Iopetalum group to be derived from allopolyploidization, 3) support an earlier hypothesis by confirming S. verrucosum as the maternal genome donor for the polyploid species S. demissum as well as species in the Iopetalum Group, 4) demonstrate that S. verrucosum is the closest relative to the maternal genome donor for species in ser. Longipedicellata, 5) support the close relationship between S. acaule and diploid species from series Megistacroloba and Tuberosa, and 6) show the North and Central American B genome species to be well distinguished from the A genome species of South America.
A comprehensive taxonomic revision of the genus Notobubon (Apiaceae) is presented. Twelve woody evergreen species are recognised, all (with the exception of N. laevigatum) endemic to the Cape Floristic Region of South Africa. The taxonomy of these prominent, though poorly collected, species has until now been problematic. They are distinguished from one another by their habit (size and branching pattern), the overall shape, size, and colour of the ultimate leaflet segments, the inflorescence structure (peduncle length, number, and length of rays in the primary umbel), the fruit morphology (fruit size, presence or absence of wings), and the fruit anatomy (symmetry of the mericarps, presence or absence of additional rib vittae, size of commissural vittae). Species relationships are assessed in the form of a cladistic analysis of 26 morphological characters, resulting in a well-resolved phylogenetic hypothesis. A comprehensive key to the species, their correct nomenclature, and typification, together with descriptions and known geographical distribution for all the species are presented and illustrated.
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