A serological contribution to the systematics of the genusLupinus (Fabaceae)

The immunological comparison of the seed reserve proteins suggests thatLupinus is a natural genus, the American and the Euro-African species belonging to the same stock. Among the Lupines of the Old World, the smooth-seeded and the rough-seeded species from two natural segregates. The genusLupinus is serologically related to the rest of theGenisteae and to the AsiaticSophoreae rather than to AmericanSophoreae andThermopsideae. The data suggest thatLupinus may have originated with the remainder of theGenisteae from primitiveSophoreae of temperate-subtropical Asia. America and the Mediterranean-African region are regarded as secondary centres of speciation.     

2C DNA variation and relationships among New World species of the genus Lupinus (Fabaceae)

The 2C DNA values in 38 species and accessions of the genus Lupinus (Fabaceae) from the New World have been analysed using flow cytometry. They are representatives of North and South American species (the Atlantic and the Andean regions). Estimated 2C DNA values ranged from 1.08 pg in L. pusillus to 2.68 pg in L. albicaulis (both from North America), that is a variation of more than 2.5-fold. The variation for North American lupins was much higher than that for South American ones. Statistical analysis of the data resulted in a grouping that showed for North American lupins some correlation with the length of life cycle. Discussion concerns some aspects of the evolution of the genus.     

Chloroplast DNA restriction site variation in theVernonieae (Asteraceae), an initial appraisal of the relationship of New and Old World taxa and the monophyly ofVernonia

Chloroplast DNA restriction site variation was examined for 35 taxa in theVernonieae and four outgroup tribes, using 17 restriction enzymes mapped for ca. 900 restriction sites per species; 139 mutations were found to be phylogenetically informative. Phylogenetic trees were constructed using Wagner and weighted parsimony, and evaluated by bootstrap and decay analyses. Relationships of Old and New World taxa indicate complex geographical relationships; there was no clear geographic separation by hemisphere. The relationships between Old and New World Vernonias found here support prior morphological analyses. The sister group to all New and most Old World taxa was composed of a small group of Old World species including yellow-flowered, trinervate-leaved species previously postulated to be basal in the tribe. The majority of both New and Old World taxa are derived from a lineage beginning with the monotypic genusStokesia, an endemic of the southeastern United States. The genusVernonia was also found to be paraphyletic within both the New and Old World. Available data do not support either the separation ofVernonia or the tribeVernonieae into geographically distinct lineages. The pattern of relationships within theVernonieae for taxa from North America, Asia, Africa, Central and South America is most similar to that of several other groups of both plants and animals with a boreotropical origin, rather than an origin in Gondwanaland. Such a pattern of distribution suggests more ancient vicariant events than are routinely postulated for theAsteraceae.     

Electrophoretic patterns of seed albumins in the Old-WorldLupinus species (Fabaceae): Variation in the 2S albumin class

PAGE and SDS-PAGE electrophoretic analysis of 31 accessions of 10 Old-WorldLupinus species, 5 smooth-seeded and 5 rough-seeded, covered total seed albumins and 2S albumins isolated by a solid-phase extraction. PAGE albumin patterns showed a distinct difference between the smooth- and rough-seeded lupins. SDS-PAGE analysis of seed albumins revealed interspecific differences, mainly due to the 2S albumins. The differences were especially marked in the smooth-seeded species. In the rough-seeded lupins the following subgroups were distinguished: (1)L. atlanticus, (2)L. cosentinii andL. digitatus, (3)L. palaestinus andL. pilosus. Evidence was provided that the 2S albumin class contains conglutin , so far classified as a globulin. The results are discussed with reference to taxonomic relationships of the Old-World lupins and characterization of the lupin seed albumin fraction.     

Comparative study of seed albumins in the Old-WorldLupinus species (Fabaceae) by reversed-phase HPLC

Seed albumins and 2S proteins isolated from the albumin fraction of 36 accessions representing 10 Old-WorldLupinus species (5 smooth- and 5 rough-seeded) were studied using reversed-phase high-performance liquid chromatography. In addition, the globulin fraction was analyzed to determine its 2S protein content. The performed separations showed the suitability of RP-HPLC technique in the analysis of variation of the seed albumin composition in lupins. In the group of rough-seeded lupins, 3 types of RP-HPLC elution profiles of albumins were distinguished: (1)L. atlanticus, (2)L. cosentinii andL. digitatus, (3)L. palaestinus andL. pilosus. All the species of this group were found to have proteins not observed in smooth-seeded species. Smooth-seeded species exhibited more abundant protein spectra, each species distinguishing by its specific RP-HPLC elution profile. It was found that 2S proteins classified as 2S albumins were responsible for the observed variation. Depending onLupinus species, the 2S albumin class consists of two to six proteins.     

Molecular phylogeny, diversification and character evolution in Lupinus (Fabaceae) with special attention to Mediterranean and African lupines

Phylogenetic relationships within the complex genus Lupinus are estimated from internal transcribed spacer (ITS) sequences of the nuclear ribosomal DNA repeat. The molecular data supports Lupinus as a distinct monophyletic group within the tribe Genisteae. Different geographical lineages are revealed within Lupinus, which are each restricted to either the Old or the New World. In the New World, the ITS data support an eastern-western geographic disjunction of the lupines and the recognition of some well-supported clades. In the Old World, almost all the previously recognized taxa are taxonomically well differentiated. The homogeneous African rough-seeded lupines, Scabrispermae, are strongly supported as a monophyletic group, which is distinct from the diverse and heterogeneous circum-Mediterranean smooth-seeded ones. The latter appear to have evolved as two lineages, in which are revealed some intersectional relationships. Also ITS data allow the assessment of the phylogenetic position of the newly discovered species, L. anatolicus (in the Old World) and L. jaimehintoniana (the Mexican tree lupin). The ITS phylogeny suggests a rapid initial radiation of the lupines subsequent to their divergence from a common ancestor. Moreover, the results indicate that the annual and perennial habits have evolved many times in Lupinus and suggest a role for generation time in affecting the evolutionary history of lupines. Data on adaptive processes and character evolution are re-examined and discussed in the light of the ITS phylogeny.These studies were supported by the research unit UMR-CNRS 6553 Ecobio – University of Rennes (France), and by the NSERC of Canada (grant to R.J. Bayer, at the University of Alberta). They are greatfully acknowledged. All the persons that have contributed in different ways to this work on Lupinus, summarized in this presentation, are greatly thanked. Particularly, we would like to mention Malika Aïnouche, Roland Greinwald, André Huon, W.K. Swiecicki, Billie L. Turner and Ludger Witte for their contributions.     

Diversification of Lupinus (Leguminosae) in the western New World: derived evolution of perennial life history and colonization of montane habitats

Previous phylogenetic studies of Lupinus (Leguminosae) based on nuclear DNA have shown that the western New World taxa form a monophyletic group representing the majority of species in the genus, with evidence for high rates of recent diversification in South America following final uplift of the Andes 2–4 million years ago (Mya). For this study, three regions of rapidly evolving non-coding chloroplast DNA (trnL intron, trnS–trnG, and trnT–trnL) were examined to estimate the timing and rates of diversification in the western New World, and to infer ancestral states for geographic range, life history, and maximum elevation. The western New World species (5.0–9.3 Mya, 0.6–1.1 spp./My) comprise a basally branching assemblage of annual plants endemic to the lower elevations of western North America, from which two species-rich clades are recently derived: (i) the western North American perennials from the Rocky Mountains, Great Basin, and Pacific Slope (0.7–2.1 Mya, 2.0–5.9 spp./My) and (ii) the predominantly perennial species from the Andes Mountains of South America and highlands of Mexico (0.8–3.4 Mya, 1.4–5.7 spp./My). Bayesian posterior predictive tests for association between life history and maximum elevation demonstrate that perennials are positively correlated with higher elevations. These results are consistent with a series of one or more recent radiations in the western New World, and indicate that rapid diversification of Lupinus coincides with the derived evolution of perennial life history, colonization of montane habitats, and range expansion from North America to South America.     

Observations on pollination inSarcostemma (Asclepiadaceae)

Field observations on pollination in New World species of the genusSarcostemma R. Br. sensuHolm are reported. Morphological and anatomical comparisons of pollinated flowers ofSarcostemma subg.Ceramanthus Kuntze (New World) andSarcostemma subg.Sarcostemma (Old World) are presented.     

Catapyrenium (Verrucariaceae) species from South America

13 species of the lichen genusCatapyrenium are reported from South America. Five species (C. analogicum, C. andicolum, C. exaratum, C. lachneoides, andC. podolepis) are described as new. A key to the species known from South America is presented. Remarks on taxonomy, ecology and distribution of the species are given.Studies on the lichen genusCatapyrenium (Verrucariaceae) III. For second part seeBreuss (1991).     

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.     

Chloroplast DNA restriction site polymorphism inGenisteae (Leguminosae) suggests a common origin for European and American lupines

Restriction site polymorphism in cpDNA of 35 legumes was studied in order to address natural relationships and geographic distribution within the tribeGenisteae. 386 sites were studied, 277 were polymorphic, 207 were informative. Phylogenetic inferences with distance and parsimony methods suggest that the American and MediterraneanLupinus species belong to a monophyletic group which arose from a single center of diversification. The data furthermore indicate thatLupinus should not be included in the tribeGenisteae since at the level of cpDNA polymorphismAnagyris foetida (tribeThermopsideae) appears more closely related to otherGenisteae thanLupinus does.     

Flavonoids as taxonomic markers in old world Lupinus species

In a leaf survey of 54 specimens of 11 Old World Lupinus species three classes of flavonoids were detected: flavones (in 82%), flavonols (in 36%) and flavone C-glycosides (in 55%). The rough-seeded species were clearly distinguished from the smooth-seeded taxa by the presence of a novel 2′-hydroxyflavone, luteolin and flavone C-glycosides as major leaf constituents and by the absence of flavonols. Within the smooth-seeded species, there are three flavonoid patterns: (a) flavonols only, L. albus; (b) flavones and flavonols, L. luteus, L. hispanicus and L. angustifolius; and (c) flavones only, L. micranthus. L. angustifolius further differed in uniquely producing diosmetin as a major leaf constituent. These divisions coincide exactly with previous groupings based on alkaloidal and morphological data. Amongst the 12 samples of L. angustifolius three chemical races were distinguished and a number of diosmetin glucoside malate esters detected. The flower flavonoid aglycone patterns of the nine Old World species surveyed differed markedly from the corresponding leaf profiles by the presence of flavones: luteolin and apigenin in eight and chrysoeriol in seven species as major constituents, while flavone C-glycosides were found only in trace amount in three species. In a leaf flavonoid survey of 13 representative New World Lupinus taxa, glycoflavones were major leaf components, a variety of methylated flavones were identified and flavonols were absent. The presence of the novel 2′-hydroxyflavone in five New World species may indicate some evolutionary link with the rough seeded taxa of the Old World.     

Phylogeny of theAsterales sensu lato based onrbcL sequences with particular reference to theGoodeniaceae

TherbcL gene of 25 taxa was sequenced and analyzed cladistically in order to define more precisely the orderAsterales s.l. and to reconstruct the phylogeny ofGoodeniaceae. The cladistic analyses show that theAsterales comprise the familiesAbrophyllaceae, Alseuosmiaceae, Argophyllaceae, Asteraceae, Calyceraceae, Campanulaceae s.l.,Donatiaceae, Goodeniaceae (includingBrunoniaceae),Menyanthaceae, Pentaphragmataceae, andStylidiaceae. Abrophyllaceae, Alseuosmiaceae, Brunoniaceae, andDonatiaceae have previously not been studied in this respect. Within theGoodeniaceae, four groups supported by therbcL data can be distinguished: the genusLechenaultia, theAnthotium-Dampiera-group, the genusBrunonia, and a group formed by the remaining genera, theScaevola-Goodenia-group.     

Comparative capitular morphology and anatomy of Coreopsis L. and bidens L. (compositae), including a review of generic boundaries

The capitular and floral morphology and anatomy ofBidens L. andCoreopsis L. were studied. All the North American species ofCoreopsis were studied. Selected species ofBidens from North and South America andCoreopsis from South America were included. The results were compared with previous observations on African species ofBidens (incl.Coreopsis). Emphasis was given to character states of the ray florets, paleae, stylearm apices, outer phyllaries, achenes, and pollen grains. Some of the character states are unique features ofCoreopsis, e.g., globular and elongately conical receptacles, deltoid outer phyllaries, truncate and indistinctly 3–5-dentate, 3–4-lobed ray florets, narrowly spathulate paleae, subulate paleae with linear-filiform upper half, hairy and apically 3-cleft paleae, truncate, convex or shallowly conical stylearm apices with the sweeping hairs limited to the area above the stigmatic surfaces and the orbicular to circular achenes. The cylindric setaceous pappus bristles so commonly encountered inBidens are unknown inCoreopsis. The pappus bristles inCoreopsis are paleaceous but similar, though thicker ones are also found in African species ofBidens (incl.Coreopsis) with winged achenes. Twin-celled hairs (setulae) with differing degrees of wall thickness are found on the achenes ofCoreopsis sect.Pseudoagarista (Mexico and South America),Coreopsis sect.Pugiopappus (California), AfricanBidens with winged achenes (e.g.,B. prestinaria, B. macroptera) and some North AmericanBidens (e.g.,B. aristosa). Similar sclerotic parenchyma make up the achenial wings of species in both genera. These may be interpreted as homologous structures, indicating the underlying similarity of these taxa and their derivation from a common ancestral stock.     

Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological andndhF sequence data

Phylogenetic relationships within the tribe Antirrhineae (Scrophulariaceae) are analysed and discussed on the basis of parsimony analyses of morphological andndhF gene sequence data. The results indicate that the tribe Antirrhineae consists of four major groups of genera, theAnarrhinum clade, theGambelia clade, theMaurandya clade, and theAntirrhinum clade. TheAnarrhinum clade, consisting of the Old World bee-pollinated generaAnarrhinum andKickxia, is sister to the rest of the tribe. TheGambelia clade consists of the New World generaGambelia andGalvezia, which are very closely related and pollinated by hummingbirds. TheMaurandya clade consists of one subclade includingMaurandya and a number of related bee- or hummingbird-pollinated New World genera and another subclade with the Old World bee-pollinated generaAsarina andCymbalaria. TheAntirrhinum clade consists mainly of bee-pollinated Old World genera, such asAntirrhinum, Linaria, Chaenorhinum, and their segregates, but also includes the New World generaMohavea andHowelliella, of which the latter is known to be partly pollinated by hummingbirds. It is concluded that hummingbirdpollination has evolved independently within Antirrhineae at least three times from bee-pollinated ancestors.     

Phylogeny and biosystematics ofPseudomonotes (Dipterocarpaceae) based on molecular and morphological data

The placement of a recently discovered South American monotypic genus,Pseudomonotes tropenbosii, in subfam.Monotoideae (Dipterocarpaceae) extends the geographical range of the subfamily from Africa to the Neotropics. Although morphological and anatomical evidence suggest similarities betweenPseudomonotes andMonotes, the close alliance of these two genera was questionable due to their disjunct distribution and a lack of phylogenetic analysis. In the present study, we reconstructed the phylogeny ofPseudomonotes and other putatively related taxa usingrbcL sequence data. The analysis ofrbcL sequences of 20 taxa belonging to 15 genera and eight families recovered a single most parsimonious tree. The genusSarcolaena (Sarcolaenaceae) formed a clade sister to the monophyleticDipterocarpaceae clade.Monotes andPseudomonotes formed a strongly supported group, sister to the monophyletic clade withPakaraimaea and the remaining Asiatic dipterocarp species studied. The study strongly supports the placement ofPseudomonotes within subfam.Monotoideae of theDipterocarpaceae.     

Magnoliid reproductive organs from the Cenomanian-Turonian of north-western Kazakhstan:Magnoliaceae andIlliciaceae

Cenomanian-Turonian sediments from the Sarbay locality in north-western Kazakhstan have yielded a rich assemblage of plant fossils including well preserved flowers, fruits, and seeds of angiosperms. This work describes fossil seeds assigned to theMagnoliaceae and theIlliciaceae. Three new species of the extinct magnoliaceous genusLiriodendroidea, L. asiatica, L. costata, andL. tenuitesta, are established and new information on the previously described species,L. alata, is provided. TheLiriodendroidea seeds are closely similar to seeds of extantLiriodendron, but are distinguished in being much smaller and winged. A new genus and species,Illiciospermum pusillum, is established based on seeds with close similarity to those of the extant genusIllicium. The seeds are small, anatropous and exotestal with outer epidermis of testa forming a palisade layer. The facets of the palisade cells have deeply undulate anticlinal walls. The micropyle area is seen on the outer integument as a transverse slit placed on a raised strophiole-like structure close to the hilum. TheIlliciospermum seeds represent the first unequivocal record of theIlliciaceae in the Cretaceous. Another seed of possible illiciaceous affinity is described as aff.Illiciospermum sp.     

Die GattungKarschia — Bindeglied zwischen bitunicaten Ascomyceten und lecanoralen Flechtenpilzen?

The genusKarschia, in the earlier sense, including saprophytes and parasites on lichens, has been thought to be a non-lichenized parallel genus of the lichen genusBuellia. Modern workers included it on the one hand inBuellia, on the other hand combined it with bitunicate ascomycetes. It is now proved thatKarschia is heterogeneous and contains but superficially similar members both of the genusBuellia of theLecanorales and of typical or masked bitunicateAscomycetes. Therefore, it can not be regarded as a link betweenLecanorales andDothideales. The type species ofKarschia belongs to theDothideales.

     

Phylogenetic analysis ofUlmaceae

A phylogenetic analysis of theUlmaceae, Cannabaceae, Barbeyaceae, andBroussonetia of theMoraceae produced nine equally parsimonious trees with 127 steps. TheUlmoideae (Ulmaceae, sensuGrudzinskaya) are a monophyletic group and distinct from theCeltidoideae. The genusAmpelocera occupies an isolated taxonomic position among the celtidoids. The similarity ofAmpelocera to the fossil celtidoid flowerEoceltis of North America suggests thatAmpelocera posesses an archaic suite of characters, and occupies a primitive position among the celtidoids, theCannabaceae and theMoraceae. The relationships among the other celtidoid taxa,Cannabaceae, andBroussonetia are problematic. TheCannabaceae andBroussonetia of theMoraceae are nested within the celtidoids suggesting that this is a paraphyletic group. The close, but unresolved, relationship of the celtidoids to theMoraceae andCannabaceae observed in this analysis, and the appearance of the celtidoids in the fossil record prior to the ulmoids suggests that the evolutionary relationship of the ulmoids and celtidoids may be more distant than current taxonomic treatments reflect.     

Chromosomal evolution inEpilobium sect.Epilobium (Onagraceae)

WithinEpilobium sect.Epilobium, a cytological analysis of 121 experimental hybrids, involving 40 species, indicates the presence of a widespread BB chromosome arrangement in Eurasia, Africa, and Australasia, as well as in North and South America less commonly. The AA chromosome arrangement, which differs from BB by one reciprocal translocation, occurs in North America, South America, and in at least three European species. The CC arrangement, which differs from AA by two reciprocal translocations, characterizes theAlpinae, a circumboreal group. Distinctive or only partly worked out chromosome arrangements occur in the EuropeanE. duriaei andE. nutans and in the North AmericanE. luteum, E. obcordatum, E. oregonense, andE. rigidum. With earlier results, the chromosome arrangements of some 65 of the estimated 185 species of the section have been established fully or partly.