Taxonomy and phylogeny of Puccinia lagenophorae: a study using rDNA sequence data, morphological and host range features

Puccinia lagenophorae is a rust fungus originating from Australasia which has spread throughout the world. A phylogenetic analysis of taxa related to this species was performed using rDNA (LSU, ITS) sequence data. The analyses revealed a well-supported cluster including all specimens of P. lagenophorae. By evaluating morphological and sequence data, the species is taxonomically re-defined and a list of synonyms is provided. Puccinia distincta on Bellis perennis, a species recently separated from P. lagenophorae, P. saccardoi, a species on the Goodeniaceae, and P. byliana, a species so far only known from South Africa, are reduced to synonymy in P. lagenophorae, as are several other species. Our analysis indicates that P. lagenophorae is likely not derived from the northern hemisphere species P. obscura, but from a species from Australia host-alternating between Asteraceae (aecial host) and Cyperaceae/Juncaceae (telial host). Another related species, P. stylidii (on Stylidium sp., Stylidiaceae) may have been derived from the same parental species as P. lagenophorae. From ontogenetical and morphological studies, the presence of pycnia could not be confirmed in the life cycle of this species, and the width of the pedicel of teliospores at the point of attachment was found to be highly variable and not a taxonomic character. The number of known host species is approximately 150, including 41 new host plants recorded herein.     

Putative natural hybrid between Puccinia lagenophorae and an unknown rust fungus on Senecio madagascariensis in KwaZulu-Natal,South Africa

Several specimens of an aecial rust fungus were collected on Senecio madagascariensis during a field survey carried out in KwaZulu-Natal, South Africa. As telia were not present in the specimens collected, DNA sequence analyses were undertaken to determine the identity of the rust species. ITS and β-tub1 sequencing confirmed that one of the isolates recovered is Puccinia lagenophorae sensu lato. On the other hand, sequencing and RFLP analysis revealed the presence of two divergent copies of ITS and β-tub1 in all the other six isolates investigated. In both phylogenetic trees, one copy of the gene region grouped within a well supported clade with sequences of P. lagenophorae accessions from different geographical origins and hosts, and the Australian rusts Puccinia saccardoi and Puccinia stylidii. The other copy of these gene regions grouped within a separate clade comprising European accessions of Puccinia dioicae (ITS) and Uromyces sommerfeltii (β-tub1) that occur on Asteraceae hosts. Multiple copies of these gene regions were not observed in Australian isolates of P. lagenophorae. Our study provides some evidence that an interspecific hybrid rust fungus, with P. lagenophorae as one of its parents, may occur on S. madagascariensis in South Africa. The identity of the other parent remains unknown.     

Morphology and phylogenetic interrelationships of the Asteraceae,Calyceraceae, Campanulaceae,Goodeniaceae, and related families (Asterales)

In search for the sister group of the Asteraceae, morphological evidence was assembled for investigating the relationship between the Asteraceae and those families most frequently considered to be their closest relatives, in particular the Calyceraceae, Campanulaceae (along with the frequently included Lobeliaceae, Cyphiaceae, Cyphocarpaceae, and Nemacladaceae), and Goodeniaceae (and the sometimes included Brunoniaceae). Several other families that have been associated with this group of families, the “Asterales-Campanulales-complex,” were also considered: Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, Menyanthaceae, and Argophyllaceae. In order to delineate the complex more precisely, another eight putatively related families were also included in the analysis. Cladistic parsimony analysis of 46 morphological and chemical characters for the 23 families was undertaken. Stability of the branches was estimated by the number of extra steps necessary to lose the group, as well as by the number of reweighted extra steps (using rescaled consistency indices) necessary to lose the group (a new approach). The results indicate that there is a monophyletic group of 14 families comprising those of the Asterales-Campanulales-complex as well as Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, and Menyanthaceae; this group is recognized as the order Asterales. Within the order, the Asteraceae, Calyceraceae, Brunoniaceae, and Goodeniaceae form one comparatively well-supported clade and the five families of the Campanulaceae sensu lato form another well-supported clade.     

Phylogenetic positions of rust fungi parasitic on ferns: Evidence from 18S rDNA sequence analysis

Molecular phylogenetic analyses of fern rusts were carried out based on 18S rDNA sequences. We sequenced the 18S rDNAs of fern rusts (Hyalopsora polypodii andUredinopsis intermedia) and non-fern rusts (Aecidium epimedii, Coleosporium asterum, Ochropsora kraunhiae, Puccinia suzutake andPhysopella ampelopsidis) and analyzed their phylogenetic relationships with other members of the Urediniomycetes. Our bootstrapped neighbor-joining tree obtained from these analyses showed that rust fungi were apparently monophyletic at high confidence level (100% bootstrap confidence). In this molecular phylogenetic tree, the two fern rusts did not occupy the basal position within the rust fungal lineage and did not form a monophyletic lineage. Two species of the Cronartiaceae (Peridermium harknessii, Cronartium ribicola) and one species of the Coleosporiaceae (Coleosporium asterum) grouped with the fern rusts. Therefore, our results suggested that the two fern rusts were not primitive. On the other hand,Mixia osmundae, which is parasitic on the primitive fernOsmunda, was phylogenetically far from the fern rusts.     

Complete chloroplast genome sequences of Praxelis (Eupatorium catarium Veldkamp), an important invasive species

Praxelis (Eupatorium catarium Veldkamp) is a new hazardous invasive plant species that has caused serious economic losses and environmental damage in the Northern hemisphere tropical and subtropical regions. Although previous studies focused on detecting the biological characteristics of this plant to prevent its expansion, little effort has been made to understand the impact of Praxelis on the ecosystem in an evolutionary process. The genetic information of Praxelis is required for further phylogenetic identification and evolutionary studies. Here, we report the complete Praxelis chloroplast (cp) genome sequence. The Praxelis chloroplast genome is 151,410 bp in length including a small single-copy region (18,547 bp) and a large single-copy region (85,311 bp) separated by a pair of inverted repeats (IRs; 23,776 bp). The genome contains 85 unique and 18 duplicated genes in the IR region. The gene content and organization are similar to other Asteraceae tribe cp genomes. We also analyzed the whole cp genome sequence, repeat structure, codon usage, contraction of the IR and gene structure/organization features between native and invasive Asteraceae plants, in order to understand the evolution of organelle genomes between native and invasive Asteraceae. Comparative analysis identified the 14 markers containing greater than 2% parsimony-informative characters, indicating that they are potential informative markers for barcoding and phylogenetic analysis. Moreover, a sister relationship between Praxelis and seven other species in Asteraceae was found based on phylogenetic analysis of 28 protein-coding sequences. Complete cp genome information is useful for plant phylogenetic and evolutionary studies within this invasive species and also within the Asteraceae family.     

Review of the sporoderm ultrastructure of members of the Asterales

Palynomorphological characteristics of the order Asterales are discussed. Particular attention is paid to the pollen morphology of basal families of this group and to that of problematic taxa that are considered as sister groups to the group under study. Ultrastructurally similar sporoderms of several families, including (1) Asteraceae, Calyceraceae, and Goodeniaceae; (2) Campanulaceae, Phellinaceae, and Menyanthaceae; (3) Rousseaceae, Abrophyllaceae, and Columelliaceae, are described. Pollen grains of Alseuosmiaceae and Stylidiaceae show unique ultrastructural features of the exine.     

The molecular phylogeny of the white blister rust genus Pustula reveals a case of underestimated biodiversity with several undescribed species on ornamentals and crop plants

Despite their economic importance, the knowledge of the biodiversity of many plant pathogens is still fragmentary. In this study we show that this is true also for the white blister rust genus Pustula that is parasitic on several genera in the asterids, including sunflower and the gentian, Eustoma. It is revealed that several distinct species exist in Pustula, suggesting that species are mostly host genus specific. No geographic patterns were observed in the occurrence of Pustula, the host range of which includes the Araliaceae, Asteraceae, Gentianaceae, and Goodeniaceae. Evidence points to these becoming hosts as a result of jumps from the Asteraceae, with subsequent host-specific adaptation and speciation. Among the undescribed species are pathogens of economic importance, e.g. the white blister rusts of sunflower, or with still restricted geographical ranges, e.g. Pustula centaurii, which could potentially spread with international seed trade, if no quarantine restrictions are implemented.     

Petal venation in the Asterales and related orders

Petal venation in families related to the Asteraceae was studied by means of light microscopy. The group of study was delimited on the base of previously published molecular data and includes the Alseuosmiaceae, Apiaceae, Aquifoliaceae, Araliaceae, Argophyllaceae, Bruniaceae, Brunoniaceae, Calyceraceae, Campanulaceae sensu lato , Caprifoliaceae, Donatiaceae, Dipsacaceae, Escalloniaceae, Goodeniaceae, Griseliniaceae, Menyanthaceae, Pentaphragmataceae, Pittosporaceae, Sambucaceae, Sphenocleaceae, Stylidiaceae, Valerianaceae, and Viburnaceae. The Calyceraceae, Goodeniaceae, and Menyanthaceae are very similar to the Asteraceae in their petal venation. One feature common to these four families but not found in any other group is the prominent marginal veins entering the petal independently of the midvein, but meeting it at the apex of the petal. Other morphological data as well as gene sequence data suggest that these four families form a monophyletic group. The other investigated families show a wide array of venation types, but they are all very different from the Asteraceae. Typical for die Campanulaceae sensu lato is a densely reticulate pattern. The Dipsacaceae, Valerianaceae and some Caprifoliaceae are characterized by well-developed transpetal veins.     

Phylogenetic species recognition reveals host-specific lineages among poplar rust fungi

Fungal species belonging to the genus Melampsora (Basidiomycota, Pucciniales) comprise rust pathogens that alternate between Salicaceae and other plant hosts. Species delineation and identification are difficult within this group due to the paucity of observable morphological features. Several Melampsora rusts are highly host-specific and this feature has been used for identification at the species level. However, this criterion is not always reliable since different Melampsora rust species can overlap on one host but specialize on a different one. To date, two different species recognition methods are used to recognize and define species within the Melampsora genus: (i) morphological species recognition, which is based solely on morphological criteria; and (ii) ecological species recognition, which combines morphological criteria with host range to recognize and define species. In order to clarify species recognition within the Melampsora genus, we applied phylogenetic species recognition to Melampsora poplar rusts by conducting molecular phylogenetic analyses on 15 Melampsora taxa using six nuclear and mitochondrial loci. By assessing the genealogical concordance between phylogenies, we identified 12 lineages that evolved independently, corresponding to distinct phylogenetic species. All 12 lineages were concordant with host specialization, but only three belonged to strictly defined morphological species. The estimation of the species tree obtained with Bayesian concordance analysis highlighted a potential co-evolutionary history between Melampsora species and their reciprocal aecial host plants. Within the Melampsora speciation process, aecial host may have had a strong effect on ancestral evolution, whereas telial host specificity seems to have evolved more recently. The morphological characters initially used to define species boundaries in the Melampsora genus are not reflective of the evolutionary and genetic relationships among poplar rusts. In order to construct a more meaningful taxonomy, host specificity must be considered an important criterion for delineating and describing species within the genus Melampsora as previously suggested by ecological species recognition.     

Studies in the Goodeniaceae and the Brunoniaceae with a discussion of their relationship to Asteraceae and Calyceraceae

The relationships between Asteraceae, Calyceraceae, Goodeniaceae, and Brunoniaceae are discussed, and some SEM-observations of microcharacters within the two latter families are presented. Based on morphological evidence neither Goodeniaceae s.str. nor Goodeniaceae s.lat. (i.e., incl. Brunonia) appear to be the putative sister taxon of Asteraceae. A cladistic analysis demonstrates that Brunonia australis may constitute a separate, monotypic family or alternatively be deeply nested within Goodeniaceae s.lat Brunonia shares a number of potential synapomorphies with Asteraceae. Of special interest are the peculiar hairs and feathery pappus bristles which very much resemble those found within Asteraceae subfam. Barnadesioideae. However, studies with SEM disclose that these structures rather constitute autapomorphies for the taxa in question. In conclusion, Calyceraceae remain to be the best sister candidate of the sunflower family.     

Fructose oligosaccharides: Possible markers of phylogenetic relationships among dicotyledonous plant families

Stems from more than 555 species representing 440 genera of approximately 210 dicotyledonous families were surveyed for the presence of fructose oligosaccharides. The tissues from 372 species representing 312 genera and 187 families did not contain such oligosaccharides. The sugars occurred sporadically or in small amounts in members of the following families: Crossosomataceae, Lauraceae, Malpighiaceae, Cornaceae, Myrisinaceae, Melianthaceae, Greyiaceae, and Simaroubaceae. The fructosyl sucrose trisaccharide, isokestose, aswell as its fructose tetrasaccharide homolog (but not higher inulin oligosaccharides) were present in members of the families Hippocastanaceae and Limnathaceae. Stems from members of the families Clethraceae and Monotropaceae, and some members of the Polemoniaceae, contained two series of fructose oligosaccharides: one based on isokestose (the inulin series) and the other one based on kestose (the levan series). The inulin or isokestose-based series of fructose oligosaccharides, alone, was present in members of the following families: Menyanthaceae, Boraginaceae, Campanulaceae, Goodeniaceae, Stylidiaceae, Brunoniaceae, Calyceraceae and Compositae. The primary metabolites occur ubiquitously and consistently in tissues of plants of these families. We hypothesize that this conservative character represents several phylogenetic alliances among them.     

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.     

南美蟛蜞菊霜霉病病原鉴定及系统发育分析

[目的]为从天敌病原生物方面探索外来入侵植物南美蟛蜞菊的生物防治,对新发现的南美蟛蜞菊霜霉病进行病原鉴定和系统发育分析。[方法]在广东省广州市对南美蟛蜞菊霜霉病的发生及危害情况进行调查,并通过病害症状识别、病原显微形态记录与比较、病原菌及其近似种ITS序列结构比较、LSU序列和ITS序列系统发育分析,对南美蟛蜞菊霜霉病病原进行鉴定和系统发育分析。[结果]南美蟛蜞菊霜霉病在广州零星发生,但该病害在华南农业大学校园内发生较严重,发病率达50%~70%,病情指数为30~35。经鉴定,其病原菌为南美蟛蜞菊单轴霉,是国内一新记录种。基于病原菌LSU序列和ITS序列的系统发育分析显示,侵染菊科植物的单轴霉属菌种聚在一个分枝,亲缘关系密切,与侵染其他不同科寄主植物的单轴霉亲缘关系较远。ITS序列结构比较显示,寄生于菊科向日葵族植物的单轴霉属菌种的ITS2区包含多个重复序列,不同菌种间的ITS2区重复序列相似度不同,说明侵染菊科向日葵簇植物的单轴霉属菌物可细分成多个种,而不是只有向日葵单轴霉。[结论]广州发生的南美蟛蜞菊霜霉病是该寄主上首次正式报道的病害,鉴定的病原菌也是国内一新记录种;寄生在菊科植物上的单轴霉属种类不尽相同,但亲缘关系紧密。     

Melampsora rust species on biomass willows in central and north-eastern Germany

Melampsora willow rusts are the most important fungal pathogens in short rotation coppices of biomass willows. In the past, breeding programmes for rust resistant biomass willows concentrated on the distinction of races within the forma specialis Melampsora larici-epitea f. sp. larici-epitea typica that colonized Salix viminalis and related clones. In a new breeding program that is based on a wider range of willow species it is necessary to identify further Melampsora species and formae specialis that are pathogens of willow species other than S. viminalis. Therefore, three stock collections with Salix daphnoides, Salix purpurea, and other shrub willow species (including S. viminalis) species were sampled in north-eastern Germany. A fourth stock collection in central Germany contributed rusts of tree willows (Salix fragilis and Salix alba) and the large shrub Salix caprea. Out of 156 rust samples, 149 were successfully sequenced for ITS rDNA. A phylogenetic analysis combining Neighbour-Joining, Maximum-Likelihood and Bayesian analysis revealed six species: Melampsora ribesii-purpureae, Melampsora allii-salicis-albae, Melampsora sp. aff. allii-fragilis, Melampsora larici-pentandrae, Melampsora larici-caprearum, and Melampsora larici-epitea. The first four species were found exclusively on the expected hosts. Melampsora larici-caprearum had a wider host range comprising S. caprea and S. viminalis hybrids. Melampsora larici-epitea can be further differentiated into two formae speciales. The forma specialis larici-epitea typica (59 samples) colonized Salix viminalis clones, Salix purpurea, Salix×dasyclados, and Salix×aquatica. In contrast to this relatively broad host range, f. sp. larici-daphnoides (65 samples) was found exclusively on Salix daphnoides. With the distinction and identification of the rust species/formae speciales it is now possible to test for race-specific resistances in a more targeted manner within the determined pairings of rust and willow species.     

The genus Uroleucon (Hemiptera: Aphididae) in the Korean Peninsula,with descriptions of two new species

The aphid genus Uroleucon Mordvilko (Hemiptera: Aphididae) is recognized from the Korean Peninsula. A total of twenty species of Korean Uroleucon are confirmed including two new species: U. (Uromelan) chrysanthemicola sp. nov., on Chrysanthemum sp. (Asteraceae) from Mt. Oseo, Korea and U. (Uroleucon) jejuni sp. nov., on Echinops setifer Iljin (Asteraceae) from Jeju Island, Korea. Two new species are described with biometric measurements, illustrations, and identification keys to species on host plants.LSID urn:lsid:zoobank.org:pub:ADBBDD99-6A12-4854-B559-64D5EF1B9CB0.     

Morphological and molecular characterisation of Puccinia lagenophorae, now present in central North America

In June 2004, a rust fungus not previously reported for Oklahoma was found occurring naturally on the weed, common groundsel, Senecio vulgaris, in pots in a commercial container nursery in northeastern Oklahoma. Host symptoms and morphology of teliospores and aeciospores of the fungus were consistent with those of Puccinia lagenophorae, a recent introduction into North America that has, as yet, been reported only on the East and West Coasts of the USA. This is the first report of the rust in central regions of North America. The rust is believed to be native to Australia and New Zealand and subsequently reported in most continents on numerous species and genera of the Asteraceae. Some authors in Europe consider the rust on Bellis as different from the one on Senecio, naming it Puccinia distincta. Our ITS1‐5.8S‐ITS2 ribosomal DNA sequence data, however, show homology with P. distincta sequences from Europe, indicating there is only one morphologically‐variable polyphagous species. Presumably, the rust was introduced into Oklahoma on infected plants from the West Coast of the USA, the source of most plant material in the commercial nursery where it was found. The rust is potentially important on several ornamental Asteraceae in North America as it has become in Europe, where it spreads rapidly throughout that continent following its introduction there in the early 1960s.     

Are the microcyclic rust species Puccinia melampodii and Puccinia xanthii conspecific?

The taxonomic relationship between two microcyclic rust species, Puccinia melampodii and Puccinia xanthii, recorded from a number of Asteraceae hosts, was explored by comparing teliospore morphology, host specificity and sequence data. Teliospore morphology varied between and within individual rust accessions but, in general, teliospores of P. xanthii were larger than those of P. melampodii. Field observations and cross-inoculation greenhouse studies showed that P. melampodii accessions were highly specific to their original host species and unable to establish compatible interactions with related Asteraceae species. Though small amounts of genetic variation were detected between P. melampodii and P. xanthii accessions, the ITS and TEF phylogenetic analyses comprising several other rust species grouped these accessions within a well supported clade. Our data indicate that despite differences between P. xanthii and P. melampodii accessions these rusts cannot readily be separated at the species level, supporting earlier taxonomic conclusions and the recent proposal to group them into a morphospecies. Based on host-specificity data, the name Puccinia xanthii var. parthenii-hysterophorae is proposed to accommodate all Mesoamerican records of P. melampodii associated with the host Parthenium hysterophorus.     

Trait convergence and diversification arising from a complex evolutionary history in Hawaiian species of <Emphasis Type="Italic">Scaevola</Emphasis>

Species variation in functional traits may reflect diversification relating to convergence and/or divergence depending on environmental pressures and phylogenetic history. We tested trait-environment relationships and their basis in finer-scale evolutionary processes among nine extant Hawaiian species of Scaevola L. (Goodeniaceae), a taxon with a complex history of three independent colonizations by different phylogenetic lineages, parallel ecological specialization, and homoploid hybridization events in Hawai‘i. Using a wild population for each species, we evaluated traits related to plant function (morphology, leaf and wood anatomy, nutrient and carbon isotope composition). Hawaiian Scaevola species were distributed across coastal, dry forest and wet forest environments; multivariate environmental analysis using abiotic and biotic factors further showed that species from distantly related lineages inhabited similar environments. Many traits correlated with environment (based on the multivariate environmental analysis), considering both distantly related species and more closely related species. Scaevola species within shared habitats generally showed trait convergence across distantly related lineages, particularly among wet forest species. Furthermore, trait diversification through divergence was extensive among closely related Scaevola species that radiated into novel environments, especially in plant morphology and traits affecting water relations. Homoploid hybrid-origin species were “intermediate” compared to their ancestral parent species, and possessed trait combinations relevant for their current habitat. The diversity in functional traits reflected strong influences of both ecology and evolutionary history in native Hawaiian Scaevola species, and trait correspondence with environment was due to the combination of multiple processes within the taxon: trait pre-adaptation and filtering, evolutionary convergence, divergence, and hybridization.