Biosystematics of two sympatric species ofEucharis (Amaryllidaceae)

Eucharis candida andE. formosa are two often sympatric species of bulbous geophytes restricted to neotropical rain forest understory. The species are most common in eastern Ecuador, and are the only two east Ecuadorean species of the genus found north of the Pastaza valley. Data from phenetic, karyotypic, and preliminary isozyme electrophoretic analyses of both species are represented. The species are distinguishable phenetically and karyologically, but isozyme-based relationships are more complex. Phenetic resolution of the isozyme phenotypes supports recognition of two species in Ecuador. A Peruvian isolate ofE. formosa, though not morphologically distinct, shows both allozyme and chromosomal divergence from Ecuadorean populations. Cladistic relationships based on overall allozyme data do not support species distinction, but a novel electrophoretic phenotype for glutathione reductase is shared only by individuals ofE. candida. An apparent geographic component within the monophyletic groups resolved in the cladogram suggests that some degree of gene flow between these two species has been maintained without the complete loss of morpholgoical species identity. This may have been mediated either by artificial population structures due to a probable long history of cultivation, or via Pleistocene refugia effects. Both species may have originated in eastern Ecuador from a common ancestral population which has since radiated outward, perhaps several times.     

Highly repeated DNA analysis and systematics of the Lemuridae,a family of Malagasy prosimians

Hybridization of highly repeated DNA sequences ofEulemur fulvus mayottensis, Lemur catta, andVarecia has been performed on blots of different species of Lemuridae (L. catta, Hapalemur griseus, Varecia variegata variegata, V. v. rubra, E. macaco macaco, E. coronatus, E. mongoz, andE. rubriventer). The probe ofE. fulvus only hybridized with the differentEulemur species, whereas that ofVarecia hybridized with the two subspecies ofVarecia and that ofL. catta with bothL. catta andHapalemur. These results were used to confirm the classification ofVarecia in a separate genus and to review the separation of theL. catta/Hapalemur group from the other species ofEulemur. Comparison of the migration patterns from DNA fragments of these different species has been used to propose a cladogram of the differentEulemur species.     

A phylogenetic analysis of <Emphasis Type="Italic">Leucothoe</Emphasis> s.l. (Ericaceae; tribe Gaultherieae) based on phenotypic characters

Phylogenetic relationships within the genus Leucothoe s.l. (including all eight species) and related taxa of the Gaultherieae, Andromedeae, and Vaccinieae were investigated by a cladistic analysis based on phenotypic (external morphology, anatomy, chromosome number, and secondary chemistry) characters. The parsimony analysis resulted in two most parsimonious trees, both very similar, which show Leucothoe s.l. to be polyphyletic, with its species distributed among three distinct clades. Our results indicate that L. racemosa and L. recurva form a strongly supported clade, which is sister to Chamaedaphne calyculata, and these three species are probably the sister-group of the wintergreen clade (consisting of Gaultheria and Diplycosia). Leucothoe axillaris, L. fontanesiana, L. davisiae, L. griffithiana, and L. keiskei, consistently form a monophyletic group corresponding to Leucothoe s.s., which is probably sister to the remaining members of the tribe Gaultherieae. Leucothoe grayana, the final species traditionally placed in the genus, belongs to neither of these clades and may be sister to Andromeda. Phenotypic characters provide no support for the monophyly of Leucothoe, instead suggesting that it is polyphyletic, in agreement with preliminary DNA-based analyses. Thus, we redefine the genus Leucothoe, placing its species into three genera: 1) Eubotrys (the E. racemosa + E. recurva clade), 2) Leucothoe s.s. (the L. axillaris + L. fontanesiana + L. davisiae + L. griffithiana + L. keiskei clade), and 3) Eubotryoides (containing only E. grayana).     

Molecular systematics of the Old World Astragalus (Fabaceae) as inferred from nrDNA ITS sequence data

This study represents a nuclear rDNA ITS-based phylogenetic analyses of a greater sampling of the Old WorldAstragalus compared to our previous work (212 vs. 134 taxa). Phylogenetic relationships among 212 species (213 accessions) of the Old WorldAstragalus, including newly segregated monotypic genusPodlechiella, the two aneuploid New WorldAstragalus, and five related genera, were inferred from analyses of nuclear rDNA ITS sequences using maximum parsimony. A total of 658 nucleotide sites and four binary characters for indels were analyzed. The results of phylogenetic analyses suggest sect.Phyllolobium, comprising mostly the Chinese species, is placed outside of the so-calledAstragalus s. str. and is a well-supported monophyletic group. The monotypic annual segregate genusThlaspidium (≡Astragalus sect.Thlaspidium, A. thlaspi), is clearly nested withinAstragalus s. str. Among the many sections analyzed here, only sects.Cenanthrum, Caraganella, Eremophysa, Incani, Laxiflori, andLotidium are strongly supported as monophyletic. Our analysis, in agreement with previous studies, shows that the North American euploidAstragalus species are scattered throughout the Old World groups of the genus.     

Phylogenetic analysis of Leucojum and Galanthus (Amaryllidaceae) based on plastid matK and nuclear ribosomal spacer (ITS) DNA sequences and morphology

Phylogenetic analyses of the monocotyledonous genera Leucojum and Galanthus (Amaryllidaceae, Asparagales), using plastid (trnL-F and matK) and largely non-coding nuclear ribosomal (ITS) DNA sequences show the two to be closely related to Lapiedra, Narcissus, Vagaria, Pancratium and Sternbergia. We compare the results obtained with a combined parsimony analysis of these nucleotide sequences with that of a matrix of morphological characters. The sampling included all species of Leucojum and most species of Galanthus (representing all series and subseries of the genus) and used as outgroup the above mentioned genera of Amaryllidaceae shown to be close relatives. The plastid, nuclear and morphological data were analysed independently and in combination, showing that the boundaries between the two genera are not appropriate. Galanthus is monophyletic but embedded in Leucojum. On the basis of chromosome numbers and floral characters Leucojum has been previously divided into four subgenera, which have been accepted as genera by some authors. In our phylogenetic analyses (separate as well as combined), Leucojum species are separated in two primary clades corresponding to L. subgenera Ruminia + Acis and L. Leucojum + Aerosperma. The taxonomic implications of this pattern are discussed, and an alternative classification is proposed. Finally, biogeographic relationships of species of both Leucojum and Galanthus are discussed, emphasising the possible origin of the narrowly distributed taxa of Leucojum relative to the widespread species.     

The genusAntennaria (Asteraceae: Inuleae) in western North America: Morphometric analysis ofAntennaria alborosea,A. corymbosa,A. marginata,A. microphylla,A. parvifolia,A. rosea,andA. umbrinella

Multivariate analysis of vegetative and reproductive characters was used to examine morphological relatedness amongAntennaria alborosea A. E. Porsild,A. corymbosa E. Nels.,A. marginata Greene,A. microphylla Rydb.,A. parvifolia Nutt.,A. rosea Greene, andA. umbrinella Rydb. Both pistillate and staminate plants were examined. Some of the characters examined were variable in one species, but stable in another (i.e., presence or absence of papillae on the achenes). Our analyses indicate that the seven species are morphologically distinct. It is hypothesized that theA. rosea agamic complex arose through hybridization amongA. corymbosa, A. microphylla, A. umbrinella, and possiblyA. dioica (L.)Gaertn. However, hybridization between the three former species and others, as well as their subsequent morphological responses to different environmental conditions causes confusion in recognizing the taxa.Antennaria angustifolia Rydb.,A. arida E. Nels.,A. confinis Greene,A. scariosa E. Nels.,A. foliacea humilis Rydb.,A. concinna E. Nels., andA. viscidula E. Nels. are considered to represent F 1 hybrids.     

Systematic analysis of Dendrobium Swartz section Dendrocoryne in the Australian region

Dendrobium sect. Dendrocoryne is a difficult taxonomic group of allied species and species complexes, occurring on the eastern Australian coastline, Lord Howe Island and New Caledonia. Significant morphological characters were assessed from all key areas of the distribution range. Patterns of variation between species, varieties and hybrids were investigated by principal coordinate analysis. The analyses affirm 12 species in the sect. Dendrocoryne described by Schlechter (1912, 1982) and Dockrill (1969, 1992), including a cluster of small statured D. gracilicaule, D. adae, D. fleckeri, and D. finniganense, and three variable species complexes – D. speciosum, D. kingianum and D. tetragonum. Recently described D. finniganense and D. callitrophilum are interpreted as members of the section. Phylogenetic relationships are presented based on parsimony analysis of 31 morphological characters. Analyses do not support proposals to recognize new genera for D. tetragonum, D. callitrophilum and D. aemulum, or to create new species within D. tetragonum, D. jonesii, D. speciosum and D. kingianum. The phylogeny indicates that rainforest taxa are earlier lineages and xerophytic taxa are more derived.     

A phenetic study of Cassia sensu lato (Leguminosae-Caesalpinioideae: Cassieae: Cassiinae) in Thailand

Cassia L. sensu lato, a large heterogeneous genus of flowering plants, occurs naturally in the tropics around the world. Recent works based on floral morphology have supported a division of this genus into three genera, namely Cassia L. s. str., Chamaecrista Moench and Senna Mill. In order to investigate this new classification, 508 specimens of 18 taxa of the genus Cassia s.l. grown in Thailand were analyzed using cluster analysis and canonical discriminant analysis. The total 32 vegetative and reproductive morphological characters were employed in these analyses. In cluster analysis, Cassia s.l. can be separated into four groups, respectively viz. Chamaecrista, Senna alata, Senna and Cassia s /str. The four-cluster grouping is discussed. From a canonical discriminant analysis using the four-cluster grouping as a priori groups, it can be concluded that Cassia s. str., Senna, and Chamaecrista are indeed distinct taxa. The three most important characters that separate the three genera are filament length, fruit length, and ovary stalk length. These quantitative characters, together with some qualitative characters, were useful in constructing an identification key to these genera. Among the three genera, it was also found that Senna is rather a heterogeneous taxon. The difference between the studied species was discussed.     

Nutritional physiology of type isolates of currently accepted species ofExophiala andPhaeococcomyces

Nutritional physiological and tolerance tests were performed for all type strains of species currently classified in the black yeast generaExophiala andPhaeococcomyces, including some additional type strains of taxa recently reidentified asExophiala species. Most describedExophiala species can be distinguished by physiological characters.Exophiala jeanselmei with its varieties, andE. castellanii should all be retained as separate taxa. The pairs of strainsMycotorula schawii/Exophiala dermatitidis, Hormodendrum negronii/Exophiala jeanselmei var.lecaniicorni andSporotrichum gougerotii/Torula bergeri were found to be conspecific. Phenetic analyses of physiological data support the identity ofPhaeococcomyces exophialae as a yeast-like synanamorph ofExophiala spinifera. The taxonomic positions of the generaNadsoniella, Phaeoannellomyces andWangiella are discussed. The generaExophiala andPhaeococcomyces are unrelated.     

Molecular variation in Galdieria sulphuraria (Galdieri) Merola and its bearing on taxonomy

Cyanidium caldarium, Cyanidioschyzon merolae and Galdieria sulphuraria are three unicellular algae characteristic, of acid thermal environments. Recently, on the basis of morphological characters, three new species of Galdieria (G. partita, G. daedala, G. maxima ) isolated from acid-thermal springs in Russia have been instituted. A selected region of rbcL and the sequence of the intergenic spacer between the rbcL and rbcS have been amplified and sequenced from different Galdieria species and strains, in order to define molecular relationship among these interesting algae. The obtained cladogram shows that Cyanidium caldarium and Cyanidioschyzon merolae form a sister group which, in turn, is in a sister group relationship with Galdieria. This last genus is divided in two clades, one of which includes G. sulphuraria accessions from Naples (Italy), California, and Yellowstone and the other one includes G. sulphuraria accessions from Java (Indonesia) and from the Russian species. These results support the status of the genus Galdieria and suggest that G. daedala, G. maxima and G. partita are three very similar strains of G. sulphuraria; the rbcL variation within Galdieria accessions has a pattern which is broadly connected to the geographial distribution. The data obtained from the intergenic rbcL-rbcS spacer partly confirm those from the rbcL analysis.     

Taxonomy and phylogeny of the tribePlucheeae (Asteraceae)

The tribePlucheeae (Benth.)A. Anderb., has been analysed cladistically by means of a computerized parsimony program (Hennig 86), using theArctotideae as outgroup. The results of the analysis are presented in a consensus tree and one cladogram. Four major monophyletic subgroups can be recognized: TheColeocoma group (3 genera), thePterocaulon group (3 genera), theLaggera group (6 genera), and thePluchea group (12 genera). All recognized genera are described and most genera are supplied with taxonomical notes including comments on their taxonomic status. Genera such asBlumea, Pluchea, andEpaltes are demonstrated to be unnatural assemblages.Monarrhenus andTessaria are both closely related to thePluchea complex. The old generic nameLitogyne Harv. has been taken up for one species ofEpaltes, the genusRhodogeron is reduced to a synonym ofSachsia, and the following new combinations are made;Litogyne gariepina (DC.)A. Anderb., andSachsia coronopifolia (Griseb.)A. Anderb.     

The phylogenetic utility of chloroplast and nuclear DNA markers and the phylogeny of the Rubiaceae tribe Spermacoceae

The phylogenetic utility of chloroplast (atpB-rbcL, petD, rps16, trnL-F) and nuclear (ETS, ITS) DNA regions was investigated for the tribe Spermacoceae of the coffee family (Rubiaceae). ITS was, despite often raised cautions of its utility at higher taxonomic levels, shown to provide the highest number of parsimony informative characters, in partitioned Bayesian analyses it yielded the fewest trees in the 95% credible set, it resolved the highest proportion of well resolved clades, and was the most accurate region as measured by the partition metric and the proportion of correctly resolved clades (well supported clades retrieved from a combined analysis regarded as “true”). For Hedyotis, the nuclear 5S-NTS was shown to be potentially as useful as ITS, despite its shorter sequence length. The chloroplast region being the most phylogenetically informative was the petD group II intron.We also present a phylogeny of Spermacoceae based on a Bayesian analysis of the four chloroplast regions, ITS, and ETS combined. Spermacoceae are shown to be monophyletic. Clades supported by high posterior probabilities are discussed, especially in respect to the current generic classification. Notably, Oldenlandia is polyphyletic, the two subgenera of Kohautia are not sister taxa, and Hedyotis should be treated in a narrow sense to include only Asian species.     

Molecular phylogeny of the Acre clade (Crassulaceae): Dealing with the lack of definitions for Echeveria and Sedum

The phylogenetic relationships within many clades of the Crassulaceae are still uncertain, therefore in this study attention was focused on the “Acre clade”, a group comprised of approximately 526 species in eight genera that include many Asian and Mediterranean species of Sedum and the majority of the American genera (Echeveria, Graptopetalum, Lenophyllum, Pachyphytum, Villadia, and Thompsonella). Parsimony and Bayesian analyses were conducted with 133 species based on nuclear (ETS, ITS) and chloroplast DNA regions (rpS16, matK). Our analyses retrieved four major clades within the Acre clade. Two of these were in a grade and corresponded to Asian species of Sedum, the rest corresponded to a European–Macaronesian group and to an American group. The American group included all taxa that were formerly placed in the Echeverioideae and the majority of the American Sedoideae. Our analyses support the monophyly of three genera – Lenophyllum, Thompsonella, and Pachyphytum; however, the relationships among Echeveria, Sedum and the various segregates of Sedum are largely unresolved. Our analyses represents the first broad phylogenetic framework for Acre clade, but further studies are necessary on the groups poorly represented here, such as the European and Asian species of Sedum and the Central and South American species of Echeveria.     

The vicarious differentiation of the alpineGalium serpylloides group (Rubiaceae), endemic to the W. Himalaya

Galium serpylloides agg. comprises a group of locally endemic, disjunct and vicarious taxa in the (sub)alpine zone of the W. Himalaya, from W to E:G. gymnopetalum, G. lahulense, G. serpylloides s. str.,G. saipalense, andG. nepalense. Four of these are new to science;G. serpylloides is newly typified. The main characters of the five species are compared and illustrated, their affinities and evolution are discussed, and a map of distribution is presented.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

Evolution,pollination, and systematics of the tribeNeottieae (Orchidaceae)

The various classifications of the orchid tribeNeottieae are reviewed and a new classification is proposed that divides the tribe into three subtribes,Neottiinae, Limodorinae, andCephalantherinae, based primarily on characters of the column (gynostemium). A cladistic analysis illustrates that these three subtribes are more closely related to one another than either is to any other group in subfam.Neottioideae, although there are very few apomorphic characters for the tribe. Pollination biology is also discussed showing links between breeding systems and distribution. There is also a possible role between column and labellum morphology and the emergence of a deceptive pollination syndrome from one of reward.     

Phylogenetics and evolutionary morphology of the Neotropical true bug genus Epipolops (Hemiptera: Heteroptera: Geocoridae)

Species of Epipolops Herrich‐Schaeffer (Hemiptera: Geocoridae), comprising the largest genus of Pamphantinae, are among the most bizarre true bugs because of their striking morphology. To elucidate evolutionary morphology in Epipolops, a phylogenetic analysis was performed using 17 species and 36 adult morphological characters. Two cladograms were obtained under equal and implied weight analyses, showing slight differences between them. Two new species, E. stridulatus sp.n . and E. univallensis sp.n ., are described, and E. meridionalis Pirán is resurrected from synonymy with E. frondosus Herrich‐Schaeffer. A key to the known species of Epipolops is provided. The systematic relevance of the unique characters of the genus is discussed and the sequence of character state transformations for both the anterior and posterior lateral processes of the pronotum are optimized on the cladogram obtained under implied weights. Species of Epipolops are found in the Neotropical region and the Mexican Transition Zone, with some clades and species restricted to certain South American subregions. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:CB72D5DA‐D86B‐4B91‐93A2‐88894F7120C9 .     

Phylogenetic relationships among species and genera of Lycoperdaceae based on ITS and LSU sequence data from north European taxa

Phylogenetic relationships, limits of species, and genera within Lycoperdaceae, were inferred by use of ITS and LSU nu-rDNA sequence data. Lycoperdaceae was confirmed as monophyletic, and Mycenastrum corium as a sister taxon to the ingroup. Four major clades were identified and received weak to moderate support and correspond with the genera Lycoperdon, Bovista, Calvatia, and Disciseda. The Lycoperdon clade includes species from Lycoperdon, Vascellum, Morganella, Handkea, Bovistella, and Calvatia. The structure within the Lycoperdon clade is unresolved and several clades are more or less unsupported, which suggests treating the supported Lycoperdon clade as the genus Lycoperdon. L. nigrescens and L. caudatum occur on single branches and their phylogenetic positions could not be resolved. The phylogenetic analyses identified 31 species of Lycoperdon, 11 species of Bovista, six species of Calvatia, and two species of Disciseda. In Lycoperdon three new species were recognized. A new species closely related to B. limosa is identified and discussed. A classification of Lycoperdaceae is proposed based on the results of the phylogenetic analyses. Morphological characters of species within and among identified clades are discussed.     

A phylogenetic analysis of Pisum based on morphological characters, and allozyme and RAPD markers

Cladistic analyses of 17 wild and cultivated pea taxa were performed using morphological characters, and allozyme and RAPD (random amplified polymorphic DNA) markers. Both branch-and-bound and bootstrap searches produced cladograms that confirmed the close relationships among the wild species and cultivars of Pisum proposed by a variety of systematic studies. Intraspecific rankings were supported for northern P. humile, southern P. humile, P. elatius and P. sativum, which together comprise a single-species complex. P. fulvum, while clearly the most divergent of the pea taxa, could also be assigned to the same species complex without violating the hierarchial logic of the cladogram. Its inclusion or exclusion depends on whether the level of interfertility it displays with other pea taxa or its overall morphological and chromosomal distinction are emphasized. As suggested by previous studies, northern P. humile was the most likely sister taxon to cultivated P. sativum; although, rigorous phylogenetic evaluation revealed a close genealogical affinity among P. elatius, northern P. humile and P. sativum. Despite their limited number, the 16 morphological characters and allozyme markers used precisely organized the pea taxa into established taxonomic groupings, perhaps in part reflecting the role morphology has played historically in pea classification. The RAPD data also generally supported these same groupings and provided additional information regarding the relationships among the taxa. Given that RAPDs are relatively quick and easy to use, are refractory to many environmental influences, can be generated in large numbers, and can complement traditional characters that may be limited in availability, they provide a valuable new resource for phylogenetic studies.     

A multivariate analysis of morphological variation in the Hemizygia bracteosa complex (Lamiaceae, Ocimeae)

The variation and recognition of taxa within the Hemizygia bracteosa complex was examined using multivariate techniques. Morphological characters were sampled on 197 herbarium specimens. Phenetically H. bracteosa and H. welwitschii overlap in many floral characters. However differences in leaf characters and habit maintain their morphological distinctness from each other and they are therefore upheld as species. Hemizygia ornata, synonymized under H. welwitschii in recent treatments of the genus Hemizygia, differentiates from H. welwitschii on the basis of differences in leaf and floral characters and its reinstatement at specific level is here proposed. The concept of H. linearis is revised on account of the type specimen separating from all other elements of the taxon used in the analyses. The latter form a coherent group with H. petrensis and H. canescens thus negating any attempt to give any of them specific recognition. It is proposed that they should be synonymized under the earliest name H. canescens. The specific boundary of H. petiolata is revised but its specific status maintained.     

A cladistic analysis of the tribeAstereae (Asteraceae) with notes on their evolution and subtribal classification

TheAstereae were surveyed and the genera arranged in 23 informal groups. The generic groups were used to sample representative genera for a cladistic analysis based on morphological characters. The resulting cladogram was used for discussion of evolution and subtribal classification within the tribe. The lower basic chromosome numbers x = 4, 5, 6, and 8 are interpreted as reductions from a primitive x = 9. The subtribeGrangeinae occupies a phylogenetically basal position as sister group to the rest of the tribe. This may be divided into two large groups, largely corresponding to the homochromousSolidagininae and to the heterochromousAsterinae sensu lato, i.e. including theBellidinae, Hinterhuberinae, Conyzinae, andBaccharidinae. The latter four subtribes are derived within theAsterinae, and hence reduced to synonymy. Several intercontinental relationships indicate that a geographical subdivision of the tribe should be avoided, although in our analysis most of the groups proved to be restricted to one of five major regions.