Phylogenetic relationships of Thlaspi s.l. (subtribe Thlaspidinae, Lepidieae) and allied genera based on chloroplast DNA restriction-site variation

Chloroplast DNA restriction-site variation was analyzed in 30 accessions representing 20 species from the major lineages in Thlaspi s.l. (previously described as genera by Meyer 1973, 1979) and allied genera from the subtribe Thlaspidinae (Peltaria, Teesdalia, Cochlearia, Ionopsidium, Aethionema). A total of 161 variable restriction sites were detected. Phylogenetic analyses indicated a division of Thlaspi s.l. into three groups consistent with Meyer's genera Thlaspi s. str., Microthlaspi and Noccaea/Raparia. The genus Thlaspi s.l. as currently described proved to be paraphyletic because one of its major lineages, i.e. Thlaspi s. str., appeared to be more closely related to other genera (Peltaria, Teesdalia) than to the remaining lineages of Thlaspi s.l., i.e. Noccaea/Raparia and Microthlaspi. Sequence divergence values (100 x p) between the Thlaspi s.l. lineages were similar to values between these groups and related genera (Teesdalia, Peltaria), respectively. Chloroplast DNA variation was also used to assess subtribal classification of the genera studied. The cpDNA data were inconsistent with the controversial taxonomic classifications based on morphology. The molecular data would suggest that (1) the subtribe Thlaspidinae, as traditionally described, is not monophyletic; (2) the Thlaspidinae should be reduced to a group consisting of Thlaspi s. str., Peltaria, Teesdalia, Microthlaspi, Noccaea/Raparia, and that Aethionema should be excluded from the Thlaspidinae; and (3) Cochlearia and Ionopsidium represent the subtribe Cochleariinae.     

Pachyphragma andGagria (Brassicaceae) Revisited: Molecular data indicate close relationship toThlaspi S.Str.

The systematic position ofPachyphragma macrophyllum (Brassicaceae) is unsettled. This species was either treated as a monotypic section ofThlaspi s.l. or was treated as a distinct genus.Gagria lobata apparently closely related toPachyphragma, was also given generic rank. To unravel the phylogenetic relationships ofPachyphragma andGagria, ITS sequence studies were conducted using as a reference representatives of the main lineages ofThlaspi s.l. The molecular data indicate thatGagria lobata andPachyphragma macrophyllum are closely related to members ofThlaspi s.str., an assemblage of species well separated from the otherThlaspi s.l. lineages.     

Thlaspi s.str. (Brassicaceae) versus Thlaspi s.l.: morphological and anatomical characters in the light of ITS nrDNA sequence data

 Nuclear ribosomal (ITS) DNA sequence data from representatives of eleven out of 12 genera previously included in Thlaspi sensu lato were analyzed to elucidate relationships within the former genus Thlaspi sensu lato. Sequences from Thlaspi segregates Noccaea, Masmenia, Callothlaspi, Kotschyella, Microthlaspi, and Noccidium were added to the existing data sets, and only some Thlaspi sensu lato segregates formed a monophyletic group. The recently renamed genus Pseudosempervivum, formerly part of the genus Cochlearia, has been shown to be closely related to this group as well. Microthlaspi consists of three independent lineages of which Microthlaspi umbellatum is closely related to Neurotropis. The Thlaspi s.str. taxa including the type species T. arvense, are closely related to Peltaria and Alliaria and they represent a monophyletic group. Kotschyella and Noccidium, which were also integrated into Thlaspi s.l. are not closely related to any Thlaspi lineage. Therefore, we suggest major taxonomical changes which are not in agreement with concepts based on morphological data. Our ITS based phylogeny demonstrates that subtribe Thlaspidinae comprises some additional genera such as Teesdalia, Peltaria, and Alliaria, the latter two even previously classified in other tribes than tribe Lepidieae. Received August 15, 2000 Accepted January 30, 2001     

Systematic Rearrangement of Korean <Emphasis Type="Italic">Scirpus</Emphasis> L. s.l. (Cyperaceae) as Inferred from Nuclear ITS and Chloroplast <Emphasis Type="Italic">rbcL</Emphasis> Sequences

The genus Scirpus L. s.l. (Cyperaceae) has been accepted as a polyphyletic taxon by most plant taxonomists. This genus was separated into different genera by several different authors: Scirpus s. str., Trichophorum Pers., Bolboschoenus (Asch.) Palla, Schoenoplectus (Rchb.) Palla, and Schoenoplectiella Lye. The heterogeneity of Korean Scirpus s.l. has not yet been studied. We examined 17 taxa of Korean Scirpus s.l. by morphological characters and phylogenetic analyses based on nuclear ITS and chloroplast rbcL sequences. Phylogenetic analyses using maximum parsimony, maximum likelihood, and Bayesian method provided sufficient resolution. The phylogeny revealed the polyphyly of Korean Scirpus s.l. with five distinct clades. These clades correspond to Bolboschoenus, Schoenoplectus s.str., and Schoenoplectiella, Scirpus s.str., and Trichophorum, respectively. These five genera were delimited in terms of the morphology of tuber, bract, and inflorescence. By virtue of our findings, we suggest that the 17 taxa of Korean Scirpus s.l. should be placed into five genera as follows: Bolboschoenus (two species), Schoenoplectus (three species), Schoenoplectiella (six species), Scirpus s.str. (five species), and Trichophorum (one species).     

Polyploidy, hybridization and reticulate evolution: lessons from the Brassicaceae

The family Brassicaceae is well known for its large variation in chromosome numbers, common occurrence of polyploids and many reports of interspecific gene flow. The present review summarizes studies from the past decades on polyploidization and hybridization events, recognizing them as important evolutionary forces in the family. Attention is drawn to the issue of the reconstruction of reticulated pattern of evolution resulting from allopolyploid and homoploid hybrid speciation. The research of various authors on several Brassicaceae genera is presented and discussed in the context of our current understanding of polyploid and hybrid evolution. Model species, Arabidopsis thaliana and Brassica taxa, are referred to only marginally, major focus is on a comprehensive survey of studies on about a dozen best explored non-model genera (e.g. Cardamine, Draba, Rorippa, Thlaspi). The increasing amount of genetic and genomic resources available for Brassicaceae model species provides excellent opportunities for comparative genetic and genomic studies. Future research directions and challenges are thus outlined, in order to obtain more detailed insights into the evolution of polyploid and hybrid genomes.     

A new presumably widespread species of Albugo parasitic to Strigosella spp. (Brassicaceae)

White blister rust is one of the most common diseases in Brassicaceae. Recently, molecular approaches revealed that apart from Albugo candida, several other more specialized species of the genus are causing this disease on Brassicaceae and the diversity of this group still remains largely unexplored. All newly described species have so far been sampled only from a limited geographic range, except for Albugo species which followed their invasive host from Europe to other continents. In this study we show that a previously unknown species of Albugo is causing white blister rust disease on two species of Strigosella. This species can be distinguished from other species of the genus Albugo both by its phylogenetic position and its unique oospore ornamentation that might be an adaptation to the harsh environment of the host plants. As a consequence, Albugo arenosa is described and illustrated as a new species, so far known from Strigosella africana in Iran and Spain, and Strigosella brevipes in Iran. Apart from Albugo candida and Albugo lepidii, this is the third hitherto known species of Albugo s. str. with a confirmed native distribution range of several thousand kilometres.     

Cynanchum and the Cynanchinae (Apocynaceae – Asclepiadoideae): a molecular, anatomical and latex triterpenoid study

The phylogeny of the genus Cynanchum s. str. is studied using cpDNA spacers and ITS. Morphological, anatomical and latex triterpenoid data are interpreted in light of the molecular results, and discrepancies are discussed. Vegetative characters are better indicators of relationship than floral characters, especially corona characters. The monophyly of all Malagasy species and, nested within the latter, of all stem-succulent taxa is ascertained and the genera Folotsia, Karimbolea, Platykeleba and Sarcostemma are subsumed under Cynanchum. One African species, C. galgalense, is excluded from Cynanchum.     

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.     

Blütenmorphologische und palynologische Untersuchungen an Littorella

In the monograph of the Plantaginaceae by Pilger (1937) Littorella is looked upon as a genus with three species. The indigenous Littorella uniflora (L.) Aschers. — native to the Atlantic Europe — is closely related to L. americana Fern., distributed in Eastern North America. In South America there occurs the most interesting species, L. australis Griseb., in two part-areas (in Southern Chile near Panguipulli-Lake and more southern near Lake San Martin to Seno de Ultima Esperenza and on Eastern Falkland-Island). Littorella is a perennial water-or swamp plant, growing on sandy and muddy soils. It can be living submersely throughout life without flowering and propagating by runners in this case. Its occurence, the structure of its inflorescence and ovary apparently quite different and the position of the male flower gave repeatedly rise in literature to the opinion, to seperate Littorella and Plantago far from each other. Indeed, in Littorella as well as in Plantago and the third genus Bougueria, native to the Andes of Peru and Bolivia, the leaves are arranged in rosettes, but the steams are extremely compressed. In the first enumerated genera, the axillary leafless main-axis is ending into a spherical to cylindrical ear, whereas in Littorella there is only one male flower (rarely two!) at the base of which there are sitting several female ones. These different opinions on the inflorescence of Littorella and its relationships to Plantago are discussed. In this connection, a flower- and pollenmorphological elaboration of the three species of the genus seemed to be useful. An evaluation of numerous flowers furnished the proof, that, also here, the ostensibly strict fourfoldness of the corolla is broken through and thus, indications for a possible derivation from a pentamere structure of the flower are to be found.     

Reclassification of an enigmatic downy mildew species on lovegrass (Eragrostis) to the new genus Eraphthora, with a key to the genera of the Peronosporaceae

Graminicolous downy mildews affect economical important crops like sorghum, maize and pearl millet, and also a variety of wild grasses, which pathogens have hitherto been much less studied. This also applies to an enigmatic downy mildew from Eragrostis cilianensis, which exhibits a chimeric appearance, with similarities to Basidiophora, Sclerophthora, and Sclerospora. In this study, the phylogenetic relationship of lovegrass downy mildew with respect to other genera of the Peronosporaceae was investigated and the characteristics of genera potentially associated with lovegrass downy mildew summarised from historic studies. A key based on the morphology of asexual stages is presented for all genera of downy mildews hitherto described. Based on morphological characteristics and sequence divergence, Sclerospora butleri is transferred to a new genus, Eraphthora. Lovegrass downy mildew seems to be widely distributed, as in addition to previous records from Malawi, India and Australia, we could now sample one specimen from Europe (Spilimbergo, Italy). Currently, it is unclear if Eraphthora butleri is able to infect the crop species Eragrostis tef, which might be a potential host of economic importance.     

Beiträge zur Kenntnis der Sippenstruktur des Genus Parabarium Pierre ex Spire (Apocynaceae)

The genus Parabarium is clearly separated from the other genera of the subtribe Urceolinae by different characters. The inflorescences of their taxa are heterocladical pleiothyrses, trunk synflorescences, or pleiochasiums. Further morphological and anatomical investigations deals with flower buds, leaves, pollen grains, fruits, seed, and other parts of the plants. Basing on the character analysis resulting from this the genus Parabarium is divided into two subgenera with two sections and two subsections enclosing 16 species. Two of the latter are described as news, further two varieties are set up as new ones. Two species are transferred from the genus Ecdysanthera into the genus Parabarium. Further three Parabarium species are inserted, and some more taxonomical corrections are made. — The possible relations within the genus Parabarium are made. Based on these results P. micranthum and therefor the subsection Parabarium (subgenus Parabarium) have to be regarded as the more original ones in face of P. quintaretii and the subgenus Pleiochasium. The section Burmanicum of the subgenus Parabarium has an exceptional position within the genus     

Phylogeny of the genus <Emphasis Type="Italic">Lophozia</Emphasis> (Dumort.) Dumort. s. str. inferred from nuclear and chloroplast sequences ITS1-2 and TRNL-F

Maximum parsimony and maximum likelihood phylogenetic trees were constructed for 21 taxa of Lophozia s. str. and the related genera, Schistochilopsis (5 species), Protolophozia elongata, and Obtusifolium obtusum based on combined nuclear ITS1-2 and chloroplast trnL-F DNA sequences. The trees were characterized by similar topology. It was demonstrated that the genus Lophozia s. str. was monophyletic, excluding L. sudetica, which deserved isolation into a distinct cryptic genus. The species distribution among the clades disagreed with the sections distinguished based on anatomical and morphological data. The relationships within the genus Schistochilopsis were consistent with the sectioning of the genus, based on morphological characters. Analysis of molecular data provided more precise definition of the systematic position of a number of taxa. A low level of genetic divergence of geographically distant forms was demonstrated.     

Phylogenetic relationships within the tribe Janieae (Corallinales,Rhodophyta) based on molecular and morphological data: a reappraisal of Jania1

Generic boundaries among the genera Cheilosporum, Haliptilon, and Jania—currently referred to the tribe Janieae (Corallinaceae, Corallinales, Rhodophyta)—were reassessed. Phylogenetic relationships among 42 corallinoidean taxa were determined based on 26 anatomical characters and nuclear SSU rDNA sequence data for 11 species (with two duplicate plants) referred to the tribe Corallineae and 15 species referred to the tribe Janieae (two species of Cheilosporum, seven of Haliptilon, and six of Jania, with five duplicate plants). Results from our approach were consistent with the hypothesis that the tribe Janieae is monophyletic. Our data indicate, however, that Jania and Haliptilon as currently delimited are not monophyletic, and that Cheilosporum should not be recognized as an independent genus within the Janieae. Our data resolved two well‐supported biogeographic clades for the included Janieae, an Indian‐Pacific clade and a temperate North Atlantic clade. Among anatomical characters, reproductive structures reflected the evolution of the Janieae. Based on our results, three genera, Cheilosporum, Haliptilon, and Jania, should be merged into a single genus, with Jania having nomenclatural priority. We therefore propose new combinations where necessary of some species previously included in Cheilosporum and Haliptilon.     

Morphological and molecular confirmation of Albugo resedae (Albuginales; Oomycota) as a distinct species from A. candida

The genus Albugo s.str. causes white blister rust on four families of the Brassicales, Brassicaceae, Capparaceae, Cleomaceae, and Resedaceae. Recent phylogenetic studies have revealed that several host specific lineages are present within Albugo on Brassicales, while it was also confirmed that Albugo candida has an exceptionally wide host range which extends from Brassicaceae to Cleomaceae and Capparaceae. The Albugo species infecting the Resedaceae was attributed in monographic studies as well as local floras to either A. resedae or, applying a broader species concept, to A. candida. In the present study, A. resedae specimens were morphologically and molecularly compared to the five Albugo species so far confirmed from Brassicales, A. candida, A. koreana, A. laibachii, A. lepidii, and A. voglmayrii. Both morphological differences of oospore ornamentation and phylogenetic analysis of cox2 mtDNA sequences provided evidence that A. resedae is distinct from A. candida and from the additional four species so far described from Brassicaceae. It thus seems possible that so far unknown factors restrict Albugo candida to Brassicaceae and its sister families, Cleomaceae and Capparaceae.     

Root anatomy of palms III. Ceroxyloideae,Nypoideae, Phytelephantoideae

Summary The root-anatomy of representatives of nine genera of Ceroxyloideae of the eleven genera accepted by UHL & DRANSFIELD (1987), of Nypa, the onliest genus of Nypoideae, and of all three genera of Phytelephantoideae have been studied. In Coryphoideae and Calamoideae every genus can be distinguished by the anatomy of the roots, whereas within one genus the species do not differ but slightly. This is true for the newly examined genera too and contributes both to define and identify the genera and towards an improved understanding of the relationship between genera and subfamilies. The genera of Ceroxyloideae are obviously related, they form a natural group, and even the somewhat isolated Pseudophoenix is well at home here. Both Nypa and Phytelephantoideae do not exhibit any affinities with other palms studied up to now.     

<Emphasis Type="Italic">Neolindleya</Emphasis> Kraenzl. (Orchidaceae), an enigmatic and largely overlooked autogamous genus from temperate East Asia

The Asiatic orchid species Neolindleya camtschatica (Cham.) Nevski has been omitted from the majority of relevant taxonomic surveys, including the recent Genera Orchidacearum. In most studies where the species has been included, it has been assigned to the species-rich genera Gymnadenia or Platanthera. A few morphologists recognised a new monotypic genus, Neolindleya Kraenzl., to accommodate the species. More recent molecular phylogenetic studies showed barely sufficient molecular disparity to justify generic separation of this species, but demonstrated clearly that the genus Neolindleya is only distantly related to Gymnadenia. However, the molecular phylogenies show relationships of equal strength of Neolindleya with Galearis (including Amerorchis) on the one hand and Platanthera s.l. on the other. In an attempt to better resolve the phylogenetic placement of Neolindleya, and to more clearly understand its biology and ecology, we have re-examined the morphology of this enigmatic species. Our results, based partly on SEM studies, reinforce the validity of Neolindleya as a genus, indicate a closer relationship with Galearis s.l. than with Platanthera s.l., and strongly suggest that the species became an autogam following ‘accidental’ loss of a functional bursicle. Ecologically, N. camtschatica is an opportunistic species that benefits from anthropogenic habitat disturbance.     

Beiträge zur Pollenmorphologie und Systematik der Convolvulaceen-Gattungen Maripa und Mouroucoa

Suggested by a previous communication on the occurence of two types of apertures among the pollen grains of Maripa s. l. (Convolvulaceae), 16 species of this genus were investigated palynologically. 3-zonocolpare grains mixed with 4–6-pantocolpate and 6–15-pantocolpate grains were found. The structure of the exine (infratectale bacula with or without supratectale processes) reveals further differences. The combination of these characters in the genus Maripa offers the separation into four types of pollen grains. The genus Mouroucoa is represented by its own type. These facts stress the necessity of the generical separation of the two genera. Consideration of pollen morphology yields some new aspects for subdivision of the genus Maripa.