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Phylogenetic trees of selected Clavicipitaceae. ( a ) Quartet puzzling tree : Because of unequal rate of nucleotide substitution among the positions, the matrix of maximum likelihood distances was computed using model of Tamura & Nei (1993). Gamma distributed rate with six Gamma rate categories were used as model of rate heterogeneity. Parameter alpha (0.26, S . E . 0.02) was estimated from the data set. Computation was done without clock assumption. ( b ) Maximum parsimony tree : Bremer decay indices and percentages from 500 jackknife replications are given over and under the clades, respectively. Jackknife parsimony was computed using deletion 36 %, 1000 replicas and splits > 50 %. The shortest tree length was 841 steps. The alignment contained 563 nucleotide sites from which 279 (49.6%) were constant.
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Types of asexual sporulation in 17 Claviceps species and the closely related Corallocytostroma ornicopreoides were revised in relation to the phylogeny of clavicipitaceous fungi. We observed: (1) enteroblastic conidiation from branched phialidic conidiophores typical of the genus (anamorph Sphacelia) in all species including Corallocytostroma; (2)...
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... Conidia formation on T2 medium appeared more abundant, which allowed more opportunity to observe the features of conidia and conidiogenesis. Building on previous observations of the phialidic conidiogenesis in Claviceps (Luttrell 1980;Rykard et al. 1984) and the occasional formation of the secondary conidia, Pažoutová and colleagues observed pleomorphic conidiation, i.e. primary enteroblastic (Sphacelia-like), holoblastic formation of secondary conidia (such as in C. africana) and sympodial holoblastic in C. citrina and C. zizaniae producing conidia with truncate ends (Pažoutová et al. 2004). The presence/absence of micro-conidia and the secondary conidia proved to be species-specific characteristics. ...
Ergot fungi produce toxic sclerotia that have significant impacts on the agricultural, food and pharmaceutical industries. These fungi were classified in various genera (such as Sclerotium clavus, or Spermoedia clavus) before Tulasne (1853) erected Claviceps and described three species based on variations in morphology and host ranges, viz. C. purpurea, C. microcephala, and C. nigricans. Since then, knowledge regarding the biological, clinical, and pharmaceutical perspectives of ergot has accumulated rapidly. However, a serious taxonomic examination was lacking until Langdon’s (1952) revision accepted 25 species. That was followed by intensive regional studies by Loveless in the 1960s for Rhodesia (now Zimbabwe, Africa), and Tanda in Japan (1970s – 1990s). More species names were reported and currently over 90 named taxa (species, varieties) are recorded in fungal name repositories (Mycobank and Index Fungorum). Most species were described based on morphological characteristics (sclerotia, ascomata and conidia) and host ranges. Some were characterized by their alkaloid profiles. Recently, DNA multi-locus sequence analyses (MLSA) were applied to resolve species complexes. For instance, the C. purpurea complex was separated into four species, and additional new species were recognized from South Africa and Canada. Infra- and supra-specific level genetic variations were identified via multi-locus and genomic studies. Based on five-locus phylogenies, Píchová and colleagues separated Claviceps into four sections: C. sect. Claviceps, C. sect. Citrinae, C. sect. Paspalorum and C. sect. Pusillae, for 60 species. Among these sections, several doubtful species names require clarification. Careful research on type specimens combined with molecular analyses is essential for clarifying these names.
... Mazorcas provenientes de familias S 1 seleccionadas por caracteres agronómicos fueron inoculadas con una suspensión acuosa de 100 000 macrosporas mL -1 del patógeno Sphacelia sp. obtenidas en medio T2 (Pažoutová et al., 2004). ...
... Ears from the S 1 families selected for agronomic characteristics were inoculated with an aqueous suspension of 100 000 macrospores mL -1 of the pathogen Sphacelia sp. obtained in a T2 medium (Pažoutová et al., 2004). ...
En el Colegio de Postgraduados se desarrolló la variedad sintética de maíz de grano blanco con buenas características agronómicas y resistencia al diente de caballo para zonas de transición y altiplano de México. C. gigantea ha sido reportado solo en México, sin embargo, se ha distribuido en zonas productoras de maíz en el altiplano y zonas de transición con clima fresco y alta humedad. La variedad se generó recombinando 10 líneas S1 generadas a partir de una población base de maíz con endospermo blanco y amplia base genética. La población se mejoró siguiendo un programa de selección recurrente en líneas S1. Las líneas obtenidas se seleccionaron después de inocular con macroconidios del hongo y las mejores se recombinaron para generar variedades sintéticas, entre las cuales la variedad CP-Tania 5 fue sobresaliente. Esta variedad sintética ofrece rendimiento de grano que compite con el de híbridos comerciales, la semilla es de menor costo, puede sembrarse por varios ciclos en condiciones limitantes y puede distribuirse entre agricultores.
... For example, Claviceps species are non-systemic pathogens, which generally produce enteroblastic Sphacelia-type conidia, which are associated with the honeydew production. They often possess holoblastic secondary conidia (Pažoutová et al., 2004) and infect the florets of grasses, sedges, and rushes. ...
The genus Aciculosporium (Clavicipitaceae, Hypocreales, Ascomycota) was established in 1908 for A. take, which is the causal fungus of witches’ broom of bamboo. Although the original description was valid at that time, a type specimen for A. take has not been designated. To standardize the use of this genus and species name, a neotypification and reference specimen of A. take is are proposed. Multilocus phylogenetic analyses based on DNA sequences from 28S rDNA, TEF, Tub2, Mcm7, and RPB2 revealed that A. sasicola is from a different lineage to A. take, and other material specimens from wavyleaf basket grass (Oplismenus undulatifolius) represent a distinct species proposed here as Aciculosporium oplismeni sp. nov. Chemical analysis using mass spectrometry and nuclear magnetic resonance spectroscopy, etc. showed that A. take produces four proline-containing cyclic dipeptides, which are moieties of ergot alkaloids, . although However, ergot alkaloids, lolines, peramine, indole-diterpenes, and lolitrem were not detected in the culture solvent. The presentis study offers clarification of the lineage and morphology of this genus.
... Especially the maturity of the sclerotia and the position of sections may affect the observations. Pažoutová et al. (2004) identified two types of phialidic conidia in honeydew or/and conidiomata: i.e., macroconidia of various shapes and sizes and microconidia of oval or globose shape, and observed that macroconidia stained darker than microconidia in cotton blue. Microconidia were produced by 10 out of 12 examined Claviceps species, except C. purpurea and C. citrina (Pažoutová et al. 2004). ...
... Pažoutová et al. (2004) identified two types of phialidic conidia in honeydew or/and conidiomata: i.e., macroconidia of various shapes and sizes and microconidia of oval or globose shape, and observed that macroconidia stained darker than microconidia in cotton blue. Microconidia were produced by 10 out of 12 examined Claviceps species, except C. purpurea and C. citrina (Pažoutová et al. 2004). We did observe oval or globose conidia on sclerotia produced by three of the examined species (sclerotia of C. occidentalis are not available), but they were stained at the same strength as other shapes and not particularly small, so we considered microconidia being not observed in honeydew/ young sclerotia. ...
... Claviceps. Pažoutová et al. (2004) observed holoblastic, holoblastic sympodial, and phialidic (enteroblastic) conidiation in Claviceps. The separation between holoblastic and enteroblastic ontogeny has been controversial due to intermediate patterns and lack of appropriate definitions. ...
Four ergot species (Claviceps ripicola, C. quebecensis, C. perihumidiphila, and C. occidentalis) were recognized based on analyses of DNA sequences from multiple loci, including two housekeeping genes, RNA polymerase II second largest subunit (RPB2), and translation elongation factor 1-α (TEF1-α), and a single-copy ergot alkaloid synthesis gene (easE) encoding chanoclavine I synthase oxidoreductase. Morphological features, ergot alkaloid production, and pathogenicity on five common cereal crops of each species were evaluated and presented in taxonomic descriptions. A synoptic key was also provided for identification.
... The morphology of the sexual (size, shape and colour of capitula, the stipe and ascospores) and asexual state (conidiogenesis, size, shape and colour of conidia and sclerotia) was compared among species from Fig. 2 to find additional characteristics supporting the observed clustering. The morphological data were extracted from original descriptions, from Langdon (1952) and from Pažoutová et al. (2004). Cultures grown on T2 and MEA agar (14-31 d, 24°C), the sphacelial stage or sclerotia were mounted in water and examined by DIC microscopy (Supplementary Table A.3). Additionally, specimens of C. junci, where the DNA and EA analyses failed, were included in the morphological investigation. ...
... Our analysis support the current concept of Claviceps, which includes non-systemic parasites exclusively infecting unfertilized ovaria (Pažoutová et al., 2004;Walker, 2004), albeit with unsatisfactory statistical support. The situation did not improve even after the species C. citrina, having transitional characteristics between Claviceps and Balansia, was omitted. ...
... nd data on ancestral species (e.g. C. diadema) are necessary to reconstruct the early stages of Claviceps evolution and assess the generic concept properly. Our analyses confirm the presence of four lineages inside of Claviceps (Pažoutová, 2001(Pažoutová, , 2003Lorenz et al., 2009) and the congenerity of Aciculosporium, Cepsiclava and Neoclaviceps (Pažoutová et. al., 2004;Schardl et. al., 2014a). The deep divergence is highlighted by the fact that all four lineages, described here as new sections, can be sharply distinguished based on the combination of the asexual spore type, host range, geography and secondary metabolite production. ...
The ergot, genus Claviceps, comprises approximately 60 species of specialised ovarial grass parasites famous for the production of food toxins and pharmaceutics. Although the ergot has been known for centuries, its evolution have not been resolved yet. Our approach combining multilocus phylogeny, molecular dating and the study of ecological, morphological and metabolic features shows that Claviceps originated in South America in the Palaeocene on a common ancestor of BEP (subfamilies Bambusoideae, Ehrhartoideae, Pooideae) and PACMAD (subfamilies Panicoideae, Aristidoideae, Chloridoideae, Micrairoideae, Arundinoideae, Danthonioideae) grasses. Four clades described here as sections diverged during the Paleocene and Eocene. Since Claviceps are parasitic fungi with a close relationship with their host plants, their evolution is influenced by interactions with the new hosts, either by the spread to a new continent or the radiation of the host plants. Three of the sections possess very narrow host ranges and biogeographical distributions and have relatively low toxicity. On the contrary, the section Claviceps, comprising the rye ergot, C. purpurea, is unique in all aspects. Fungi in this section of North American origin have spread all over the world and infect grasses in all subfamilies as well as sedges, and it is the only section synthesising toxic ergopeptines and secalonic acids. The evolutionary success of the Claviceps section members can be explained by high toxin presence, serving as feeding deterrents and playing a role in their protective mutualism with host plants. Closely related taxa Neoclaviceps monostipa and Cepsiclava phalaridis were combined into the genus Aciculosporium.
... Some (e.g. Claviceps species and Neoclaviceps monostipa) replace host seeds with their fruiting structures (Pažoutová, Kolarik, & Kolinska, 2004;Tudzynski & Scheffer, 2004), while others (Metarhizium species) are plant root associates and insect pathogens (Gao et al., 2011). Many form systemic (endophytic) associations throughout host shoots but fruit in a highly localized manner on leaves, nodes, buds, or inflorescences. ...
... Many form systemic (endophytic) associations throughout host shoots but fruit in a highly localized manner on leaves, nodes, buds, or inflorescences. Such systemic associations characterize Aciculosporium (Tanaka & Tanaka, 2008), Atkinsonella (Leuchtmann & Clay, 1989), Balansia (Diehl, 1950), Ephelis (Tanaka & Tanaka, 2008), Epichloe¨ (Leuchtmann, Bacon, Schardl, White, & Tadych, 2014), Heteroepichloe¨ (Tanaka & Tanaka, 2008), Myriogenospora (Glenn, Rykard, Bacon, & Hanlin, 1998), and Parepichloe¨ (White & Reddy, 1998) species and possibly also Cepsiclava phalaridis (Walker, 2004), Corallocytostroma species (Pažoutová et al., 2004), Neoclaviceps monostipa (White & Reddy, 1998), and Nigrocornus scleroticus (Ryley, 2003). Some species grow in intercellular spaces (endobiotic growth), whereas others are restricted to surfaces of, and spaces between, plant tissue layers (epibiotic growth). ...
... It seems somewhat ironic, then, that sexual Epichloe¨species fruit on immature host inflorescences and arrest their development, thereby preventing seed production from the symptomatic tillers. This phenomenon, known as "choke disease", also occurs on most or all reproductive tillers of grasses infected with Atkinsonella or Corallocytostroma species (Pažoutová et al., 2004) and on grasses and sedges (Cyperaceae) infected with certain Balansia and Ephelis species (Diehl, 1950;Tanaka & Tanaka, 2008). What is remarkable about the sexual Epichloe¨species is that, in most infected hosts, the majority of reproductive tillers are completely asymptomatic and give rise to normal seeds bearing the fungal symbiont. ...
Fungi of family Clavicipitaceae serve as models for evolution on the symbiotic continuum from pathogenic to mutualistic. Clavicipitaceous fungi associate with plants, invertebrates, and other fungi. Most plant-associated Clavicipitaceae systemically colonize shoots, but the fungal fruiting structures are localized to inflorescences, florets, buds, leaves, or nodes. Many Clavicipitaceae decrease or eliminate host seed production, but some have evolved such intimate symbioses with plant hosts that they disseminate clonally in seeds (vertical transmission) without damage or any reduction in plant fertility. In such cases, the fungi dramatically enhance host fitness by producing defensive alkaloids and through other mechanisms. To date, sequences have been assembled for 26 Clavicipitaceae representing 21 species in seven genera. These include three Claviceps species that fruit on and replace host ovaries, two Metarhizium species that parasitize insects and associate with plant roots, and 21 strains of systemic plant parasites or symbionts. Of the latter, 14 are capable of vertical transmission, and of those, 7 are strictly seed-borne mutualists in genera Epichloë and Periglandula. Alkaloid biosynthetic genes are widely distributed among these fungi. Gene clusters for ergot alkaloids and indole-diterpenes, both of which are neurotoxins in vertebrates and invertebrates, are present in members of all seven genera. The genes for anti-insect loline alkaloids and peramine have a more restricted distribution, but are present in many of the vertically transmissible Epichloë species. The availability of these genome sequences will facilitate studies of the evolution and mechanisms underlying the diversity of metabolism, host interactions, and niche adaptation of plant-associated Clavicipitaceae.
... INTA-EEA Paraná. Mayo 2014.Un examen al microscopio óptico de los tejidos de endosperma afectado mostró la presencia de abundantes conidios de S. sorghi (Figura 9), dispuestos sobre fiálides(Pažoutová et al., 2004). Sphacelia sorghi desarrollado en el endosperma. ...
... Crous and Groenewald (2005) stated that taxonomic names based solely on fungal phenotype often represent complexes of cryptic species, and not single operational units and this is especially remarkable in the phytopathogenic fungi. In the genus Claviceps, three closely related groups of species have been found to cluster around C. purpurea, C. fusiformis, and C. maximensis, each suggesting past adaptive radiations from a common ancestor (Pažoutová 2001;Pažoutová et al. 2004). The speciation process continues inside extant species: in C. purpurea, C. africana and C. fusiformis specialized populations/cryptic species have been found (Pažoutová et al. 2002a;Pažoutová et al. 2008). ...
Eight undescribed species of Claviceps were recognized from the combinations of molecular and morphological characters. The teleomorph was observed only for Claviceps setariicola. Phylogenetic affinities of the new species inside the genus were revealed by a 5.8S-ITS-28S nrDNA analysis. Claviceps chloridicola, C. tenuispora, C. setariicola and C. setariiphila are related to C. maximensis; C. truncatispora is a sister species to C. pusilla. Claviceps clavispora and C. langdonii cluster with species colonizing maize and sorghum. The position of C. loudetiae is unclear. Comparisons with herbarium specimens showed C. setariicola as a well-established species on Setaria spp. in the southern USA. C. tenuispora was recorded on Cenchrus and Pennisetum in Brazil, USA, and Zimbabwe. C. setariiphila was found on S. geniculata in Brazil. C. chloridicola, C. loudetiae and C. truncatispora occurred in African savannas on Chloris, Loudetia, and Hyparrhenia spp., respectively. C. clavispora was found on Paspalum sp. and Urochloa sp. in Mexico and C. langdonii colonized Dichanthium spp. in the southern USA and probably in Mexico. The occurrence of C. pusilla on pearl millet in the USA (Texas) is reported and the record of C. sulcata on Urochloa brizantha in Brazil is confirmed by nrDNA sequence comparison with an African herbarium specimen. No alkaloids were detected in sclerotia
and/or sphacelia of the new species.
KeywordsAscomycota–Taxonomy–Phylogeny–
Clavicipitales
–Ergot
... 4 Habitat-specific populations (races) C. purpurea differ in the type of alkaloids produced. [5][6][7] The sclerotia of population G1 contained various combinations of ergot alkaloids; all G2 produced ergosine (2) and ergocristine (1), often with small amounts of 3, and all G3 isolates belonged to a chemotype producing 1 and 3. ...
Claviceps purpurea, C. grohii, C. zizaniae, C. cyperi, and C. nigricans are closely related ergot fungi and form a monophyletic clade inside the genus Claviceps. Analysis of alkaloid content in C. nigricans sclerotia using UPLC detected ergocristine (1), ergosine (2), alpha-ergocryptine (3), and ergocristam (4). Alkaloids 1, 3, and 4 were found in the sclerotia of C. grohii. The content of 4 in the mixture of alkaloids from C. nigricans and C. grohii (over 8% and over 20%, respectively) was unusually high. Submerged shaken cultures of C. nigricans produced no alkaloids, whereas C. grohii culture formed small amounts (15 mg L (-1)) of extracellular clavines and 1. In the previously used HPLC method the ergocristam degradation product could have been obscured by the ergosine peak. Therefore sclerotia of a C. purpurea habitat-specific population G2 with the dominant production of 1 and 2 have been reanalyzed, but no 4 was detected. The phylogeny of the C. purpurea-related species group is discussed with regard to alkaloid-specific nonribosomal peptide synthetase duplication leading to the production of two main ergopeptines instead of a single product.
... take are different from those of Claviceps. Claviceps species are characterized as follows: (1) typically possessing enteroblastic conidia (anamorph Sphacelia) associated with honeydew production and often possessing holoblastic secondary conidia (Pazoutova et al. 2004); (2) infecting fl orets of grasses and replacing the host ovules with a sclerotium; and (3) forming stipitate ascostromata arising from sclerotia . Aciculosporium take is characterized as follows: (1) possessing two types of conidia: three-celled fi liform conidia with swollen ends, and two-celled holoblastic macroconidia with two dichotomously branched apical appendages that separate by breaking at the middle septum, with new germination occurring from the basal end (Tsuda et al. 1997); (2) living endophytically in vegetative shoots of bamboo (Poaceae) (Nozu and Yamamoto 1972 ); (3) forming conidiostromata and sessile ascostromata at the shoot of witches' broom twigs. ...
... Claviceps phalaridis) is a systemic endophyte of pooid grasses in southeastern Australia. It produces sclerotia in fl orets of infected plants, produces stipitate ascostromata from sclerotia, and has two anamorphs, one of which produces holoblastic conidia with apical branched appendages (Walker 2004). Pazoutova et al. (2004) showed that these Aciculosporium, Neoclaviceps, and Cepsiclava species could be grouped together, and this group is related to Claviceps on the basis of ITS sequences. In conclusion, the results of the present study also strongly suggest that Aciculosporium is phylogenetically related to Claviceps rather than Balansia. The appendaged co ...
Aciculosporium and Heteroepichloë (Clavicipitaceae) are characteristic bambusicolous fungi in east Asia. In this study, we examined their intergeneric relationships
based on the ALDH1-1 gene, which encodes a member of the aldehyde dehydrogenase family. In the clavicipitaceous fungi examined in this study,
the nucleotide sequence of the third exon of ALDH1-1 (Exon-3) is 889 bp in length and has no insertion/deletion. A phylogenetic tree based on Exon-3 indicated that the clavicipitaceous
fungi could be divided into two large groups: Cordyceps, Nomuraea, and Ustilaginoidea species formed a paraphyletic group, and the other grass biotrophic species formed a monophyletic group. This monophyletic
group was further divided into three groups with high bootstrap support: i.e., species with Neotyphodium anamorphs (e.g., Epichloë), species with Ephelis anamorphs (e.g., Heteroepichloë), and Aciculosporium-Claviceps species. We discuss the relationships among Aciculosporium, Heteroepichloë, and other clavicipitaceous fungi.
