| HOME | HELP | FEEDBACK | SUBSCRIPTIONS | ARCHIVE | SEARCH | TABLE OF CONTENTS |
Centro de Recursos Microbiológicos, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, Quinta da Torre, 2829-516 Caparica, Portugal
Artur Alves
António Correia
Centro de Biologia Celular, Departamento de Biologia, Universidade de Aveiro, Campus Universitário de Santiago, 3810-193 Aveiro, Portugal
Jordi Luque
Departament de Protecció Vegetal, Institut de Recerca i Tecnologia Agroalimentàries, Centre de Cabrils, Ctra. de Cabrils s.n., E-08348 Cabrils, Barcelona, Spain
 |
ABSTRACT |
|---|
 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONTAXONOMIC PARTLITERATURE CITED |
|---|
Botryosphaeria sarmentorum sp. nov. and B. iberica sp. nov. are described and illustrated. These two species are unusual in this genus because of their brown, 1-septate ascospores. Phylogenetic analysis based on ITS and EF1-
sequences place them within the clade containing species with Fusicoccum anamorphs. The brown, 1-septate conidia, however, do not conform to Fusicoccum. Therefore phylogenetically and morphologically the anamorphs of these two species belong in a genus distinct from any of the currently accepted anamorph genera assigned to Botryosphaeria. Through a study of the type species of Dothiorella this genus is resurrected to accommodate anamorphs of Botryosphaeria with brown, 1-septate conidia. Botryosphaeria sarmentorum is shown to be the teleomorph of Diplodia sarmentorum, which in turn is transferred to Dothiorella. Otthia quercus is transferred to Botryosphaeria as B. quercicola nom. nov.
Key words: Botryosphaeriaceae, Diplodia, Fusicoccum, ITS, molecular phylogenetics, Otthia, translation elongation factor EF1-
|
INTRODUCTION |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONTAXONOMIC PARTLITERATURE CITED |
|---|
and later emended by Saccardo (1877). Because Cesati and De Notaris (1863) did not designate a type species, von Höhnel (1909) suggested B. berengeriana De Not. as lectotype, while Theissen and Sydow (1915) suggested B. quercuum (Schwein.) Sacc. Both proposals were rejected because these species were not included in the original description. Barr (1972) proposed B. dothidea (Moug. : Fr.) Ces. & De Not. because it was one of the original species described and it conforms with Saccardo’s (1877) emendation. This proposal has been accepted generally, and Slippers et al (2004a) provided a revised description of this species based on the type specimen and fresh collections.
Botryosphaeria is well defined morphologically, and descriptions have been given by von Arx and Müller (1954), Eriksson (1981) and Sivanesan (1984). Characters of the genus are: Pseudothecia are thick-walled, multilocular or unilocular sometimes forming botryose clusters. They often are united with conidiomata on a common basal stroma and are embedded in the host, becoming partially erumpent at maturity. Septate pseudoparaphyses, often constricted at the septum, frequently are present in the centrum of immature pseudothecia, but they gradually disappear as the asci develop and mature. Asci are bitunicate with a thick endotunica, stalked or sessile, clavate with a well developed apical chamber, and they develop on a broad basal hymenial layer. Ascospores are hyaline, thin-walled, aseptate and vary from fusoid to ellipsoid to ovoid and are irregularly biseriate in the ascus. Ascospores sometimes become brown and 1-or 2-septate with age (Shoemaker 1964, Sivanesan 1984, Denman et al 2000, Alves et al 2004).
Species in Botryosphaeria were described largely on the basis of form of their ascomata and host association, and this led to a proliferation of names. Von Arx and Müller (1954) examined 183 taxa and reduced them to 11 species, with extensive synonymies under B. dothidea and B. quercuum and nine new combinations. However, because von Arx and Müller (1954) did not take into account the characters of the anamorphs and because species of Botryosphaeria are difficult to separate on the basis of teleomorph characters, these synonymies have not always been accepted (Shoemaker 1964, Sivanesan 1984, Slippers et al 2004a).
While morphology of the teleomorphs of Botryosphaeria species differ little between species, a wide range of morphologies are seen in their anamorphs. According to Denman et al (2000), anamorphs of Botryosphaeria species have been assigned to 18 coelomycete genera, among which the most common are Botryodiplodia (Sacc.) Sacc., Diplodia Fr., Dothiorella Sacc., Fusicoccum Corda, Lasiodiplodia Ellis & Everh., Macrophoma (Sacc.) Berl. & Voglino and Sphaeropsis Sacc. (Sutton 1980, Pennycook and Samuels 1985, Samuels and Singh 1986, Morgan-Jones and White 1987, Slippers et al 2004a). These genera were not delimited clearly, and several have been reduced to synonyms. Thus Sutton (1980) synonymized Macrophoma with Sphaeropsis and Denman et al (2000) suggested that Sphaeropsis and Lasiodiplodia are synonyms of Diplodia. Crous and Palm (1999) showed that Botryodiplodia is a nomen dubium and that Dothiorella should be regarded as an additional synonym of Diplodia.
Recent studies on the taxonomy of Botryosphaeria have employed molecular methods to reveal phylogenetic relationships among species ( Jacobs and Rehner 1998) and to help resolve species complexes (Smith et al 2001, Phillips et al 2002, Denman et al 2003, Alves et al 2004, Slippers et al 2004a). In these respects nucleotide sequences of the 5.8S nuclear ribosomal DNA gene and the flanking internal transcribed spacers (ITS1 and ITS2) have been used widely. Phylogenies resulting from ITS sequence analyses reveal two major clades that correspond to species with Fusicoccum or Diplodia anamorphs ( Jacobs and Rehner 1998, Zhou and Stanosz 2001). This has been interpreted as evidence for a major split in the genus into species with hyaline, fusoid, thin-walled, usually aseptate conidia (Fusicoccum) and those with hyaline or colored, thick-walled, aseptate or 1-septate, ellipsoid conidia (Diplodia) (Denman et al 2000, Zhou and Stanosz 2001). Within these two clades, several species can be distinguished on their morphology. The Fusicoccum clade has been studied in detail, and at least 11 phylogenetic species can be recognized (Smith et al 2001, Denman et al 2003, Slippers 2004a, b).
In contrast the Diplodia clade has received less attention and several taxonomic problems remain to be resolved. At present it is possible to recognize at least four phylogenetic species in this clade (Zhou and Stanosz 2001, Alves et al 2004). Van Niekerk et al (2004) described D. porosum Niekerk & Crous, which they regarded as occupying a position intermediate between Fusicoccum and Diplodia.
In this paper we describe two new species of Botryosphaeria that are unusual in having predominantly dark brown, 1-septate ascospores. Conidia of their anamorphs are thick-walled, brown and 1-septate and in this respect they resembled Diplodia sarmentorum (Fr.) Fr. (Wollenweber 1941, Laundon 1973). Because the teleomorph of D. sarmentorum is considered to be Otthia spiraeae (Fuckel) Fuckel (Booth 1958), the relationship of the two new species to Otthia was determined through a study of type specimens, and their relationship to other Botryosphaeria species was determined through a study of their morphology and ITS and EF1- sequence analysis.
|
MATERIALS AND METHODS |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONTAXONOMIC PARTLITERATURE CITED |
|---|
and incubated at 25 C under daylight fluorescent tubes. Sporulation was enhanced by placing autoclaved twigs of Populus nigra L. on the agar surface. Growth rates were determined on Difco potato-dextrose agar (PDA) plates incubated in the dark at temperatures ranging from 5 to 35 C in 5 C intervals. Representative isolates were deposited at the Centraal-bureau voor Schimmelcultures (CBS), Utrecht, The Netherlands. Specimens were lodged with the herbarium of Estação Agronómica Nacional (LISE), Oeiras, Portugal.
Type material as well as other representative specimens of the studied taxa were obtained from several herbaria, including G, IMI, K, NYS and UPS. Conidia and other structures were mounted in 100% lactic acid and digital images recorded with a Leica DFC320 camera on a Leica DMR HC microscope fitted with Nomarski differential interference-contrast optics. Measurements were made with the Leica IM500 measurement module. From measurements of 50 conidia and ascospores the mean, standard deviation and 95% confidence intervals were calculated. Dimensions are given as the 95% confidence limits with minimum and maximum dimensions in parentheses. Dimensions of other fungal structures are given as the range of at least 20 measurements.
Phylogenetic analysis.— –
The procedures described by Alves et al (2004) were used to extract genomic DNA from fungal mycelium and to amplify part of the nuclear rRNA operon using the primers ITS1 and ITS4 (White et al 1990). The primers EF1-728F and EF1-986R (Carbone and Kohn 1999) were used to amplify part of the translation elongation factor 1-alpha (EF1-) gene. PCR reactions were carried out with Taq polymerase, nucleotides and buffers supplied by MBI Fermentas (Vilnius, Lithuania), and PCR reaction mixtures were prepared according to Alves et al (2004). The amplification conditions were: initial denaturation of 5 min at 95 C, followed by 30 cycles of 30 s at 94 C, 45 s at 55 C, 1
min at 72 C, and a final extension period of 10 min at 72 C. In some cases where amplification of the EF1- region was not accomplished a second PCR was performed using as template 1 µL of the first PCR amplification.
The amplified PCR fragments were purified with the JET-QUICK PCR Purification Spin Kit (GENOMED, Löhne, Germany). Both strands of the PCR products were sequenced with the ABI PRISM® BigDyeTM Terminator Cycle Sequencing Ready Reaction Kit with AmpliTAQ DNA Polymerase (PE Applied Biosystems, Foster City, California) in a Bio-Rad iCycler Thermal Cycler. Cycle sequencing procedure was described elsewhere (Alves et al 2004).
The sequences were obtained with the ABI PRISM® 310 Genetic Analyzer (PE Applied Biosystems, Foster City, California). The sequences of the ITS (partial 18S, ITS1, 5.8S gene, ITS2 and partial 28S sequences) and partial EF1- regions were read and edited with Chromas 1.45 (http://www.technelysium.com.au/chromas.html). All sequences were checked manually, and nucleotide arrangements at ambiguous positions were clarified with both primer direction sequences. Sequences were deposited in the GenBank public database. Nucleotide sequences for both DNA regions of additional Botryosphaeria species were taken from GenBank (TABLE I).
|
sequences were aligned with MegAlign (DNASTAR Inc., Madison, Wisconsin), and manual adjustments were made where necessary. Phylogenetic analyses of sequence data were done with PAUP* version 4.0b10 (Swofford 2003). All characters were unordered and of equal weight, and alignment gaps were treated as a fifth character state. The ITS and EF1- sequences were combined and a partition homogeneity test was conducted in PAUP (Swofford 2003) to examine the possibility of a joint analysis of the two datasets. Maximum parsimony analyses were performed with the heuristic search option with 1000 random taxa addition and tree bisection and reconnection (TBR) as the branch-swapping algorithm. Branches of zero length were collapsed, and all multiple, equally parsimonious trees were saved. The robustness of the most parsimonious trees was evaluated by 1000 bootstrap replications (Hillis and Bull 1993). Other measures used were consistency index (CI), retention index (RI) and homoplasy index (HI). |
RESULTS |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONTAXONOMIC PARTLITERATURE CITED |
|---|
datasets contained 11 isolates with brown, 1-septate conidia and a further 29 isolates, either determined in this study or retrieved from GenBank, and representing 13 other Botryosphaeria species (TABLE I
). The result of a partition homogeneity test (P = 0.370) indicated that the ITS and EF1- trees reflect the same underlying phylogeny, therefore the two datasets were combined in a single analysis. Sequence alignments are available from TreeBASE (S1161), and new sequences were deposited in GenBank (TABLE I). The combined dataset contained 873 characters, of which 304 were constant, 196 variable characters were parsimony uninformative and 373 were parsimony informative. Maximum parsimony analysis resulted in eight trees (one is shown in FIG. 1). The eight trees differed only in the positions of the isolates in the terminal clades and exhibited low homoplasy as indicated by a consistency index of 0.700, a retention index of 0.888 and a homoplasy index of 0.300.
|
). The second major clade (72% bootstrap support) was resolved into 10 groups, eight of which corresponded to Botryosphaeria species with Fusicoccum anamorphs. In addition, two new species (Botryosphaeria sarmentorum A.J.L. Phillips, Alves & Luque and B. iberica A.J.L. Phillips, Alves & Luque) with anamorphs morphologically similar to D. sarmentorum fell within this clade. Isolates determined to be the anamorph of B. sarmentorum and originally recorded under the names O. spiraeae (IMI 63581b), Diplodia pruni Fuckel (CBS 165.33) and D. sarmentorum (CBS 120.41) grouped with an isolate from Malus pumila Mill. from The Netherlands (CBS 115038), while all the isolates from oaks clustered together with an isolate from M. pumila from the eastern Pyrenees ( JL 384). These two groups were supported by high bootstrap values of 
94% and differed by three base pairs in the ITS dataset and two substitutions and 13 insertions/deletions in the EF1- dataset.
Morphological characterization.— –
A total of 19 strains with brown, 1-septate conidia were isolated from Quercus spp. or Malus pumila (TABLE I). Further strains were obtained from the CABI or CBS culture collections (TABLE I). All strains, except CBS 120.41 and CBS 165.33, sporulated in culture. Conidiomata formed on Populus twigs on OA cultures were stromatic, uniloculate with a well defined ostiole. Coni-diogenesis was holoblastic with proliferation at the same level giving rise to periclinal thickenings. They rarely were seen to proliferate percurrently to form one or two indistinct annellations. Conidia in all strains were brown, thick-walled and 1-septate with both ends rounded or occasionally with a truncate base (FIGS. 3–4, 12–13, 15, 24–27). These characters fit closely with those reported for Diplodia sarmentorum (Stevens 1936, Wollenweber 1941, Booth 1958, Laundon 1973). Conidia of the isolates from oaks tended to be larger (23–26 x 9–12.5 µm) than the conidia of the two isolates from Ulmus and Malus (19–23.5 x 8.5–10.5 µm). The type specimen of Sphaeria sarmentorum Fr. in UPS (Fries Scl. Suec. No. 18) and an epitype (K104852) were examined. Conidia in these two specimens were brown, thick-walled, 1-septate and measured 20–23 x 9–11 µm. In this respect they closely resembled the isolates from hosts other than oaks. We also examined the type of Dothiorella pyrenophora Sacc. (K 54912). Pycnidia of this specimen were not fully mature, but we confirm the findings of Crous and Palm (1999) that the conidia become brown and 1-septate at an early stage.
|
|
|
|
, 19). Ascospores were brown, 1-septate, ovoid with a broadly rounded apex, and they tapered to a rounded base (FIG. 20). Pseudoparaphyses were broad and frequently septate (FIG. 18).
Because the anamorphs considered in this work were morphologically similar to D. sarmentorum and the teleomorph of this is considered to be Otthia spiraeae (Booth 1958) we examined the lectotype species of this genus (K 104853, Fuckel, Fungi Rhenani No. 975). This specimen has long, cylindrical, bitunicate asci with a thin endotunica, and each ascus bears eight, obliquely uniseriate ascospores (FIG. 37). The ascospores are brown and 1-septate, oval, with both ends rounded (FIG. 39) and the pseudoparaphyses are narrow and infrequently septate (FIG. 40). These morphological features differ markedly from those of LISE 94944 as described above, and so the name O. spiraeae cannot be applied. We also examined IMI 63581b, the specimen from which Booth (1958) isolated D. sarmentorum. The clavate asci with a thick endotunica (FIGS. 2, 9, 10), and broad, frequently septate pseudoparaphyses (FIG. 8) of the ascomycete in IMI 63581b confirm that this specimen morphologically is not a species of Otthia. It is, however, similar to LISE 94944 in the brown, 1-septate ascospores, but the size and shape of the ascospores separate these two collections. Thus, in IMI 63581b the ascospores are oval and do not taper to the base as in LISE 94944. Furthermore, asci of IMI 63581b contain 4–8 ascospores, while in LISE 94944 the asci are mostly 8-spored. These two fungi phylogenetically belong in Botryosphaeria, and morphologically they are distinct from one another and other known species in this genus. Therefore they are described here as B. sarmentorum and B. iberica.
|
|
DISCUSSION |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONTAXONOMIC PARTLITERATURE CITED |
|---|
, Sivanesan 1984, Alves et al 2004), the majority remain hyaline and aseptate. In contrast nearly all the ascospores of B. sarmentorum and B. iberica are brown and 1-septate, while hyaline, aseptate ascospores were rarely seen. Furthermore ascospore coloration of the two new species described here was more intense than is seen in other Botryosphaeria species. This suggests that ascospore coloration and septation are not useful characters at the generic level.
The brown and 1-septate ascospores of the two species described here could place them in Dothidotthia Höhn. The only differences that separate these two genera are the ascospores, which are brown and 1-or 2-septate in Dothidotthia but hyaline and aseptate in Botryosphaeria (Barr 1987, 1989). The phylogenetic analysis placed both of the new species within Botryosphaeria, thus raising the question of the validity of Dothidotthia. Morphology of IMI 63581b correlates closely with the description of Dothidotthia ramulicola (Peck) Barr as given by Barr (1987, 1989). We examined the type specimen of Sphaeria ramulicola Peck (basionym of D. ramulicola) in NYS. The brown, phragmosporous ascospores of this specimen confirmed it to be a species of Leptosphaeria Ces. & De Not. Therefore the name of L. ramulicola (Peck) Sacc. applies and the name D. ramulicola could not be applied to IMI 63581b or LISE 94944.
The anamorphs of the two species of Botryosphaeria described in this paper are also unusual in that their conidia become brown and 1-septate at an early stage in their development, often before they are released from the conidiogenous cell. Apart from Lasiodiplodia, all other Botryosphaeria species thus far subjected to phylogenetic analysis have conidia that are aseptate but sometimes they become brown and 1-septate after aging. In the phylogenetic analysis the two species described in this paper clustered within the Fusicoccum clade. However, they are morphologically distinct from Fusicoccum (Crous and Palm 1999) and cannot be accommodated in this genus. They do not belong phylogenetically in Diplodia. Clearly another anamorph genus is needed to accommodate them.
When Crous and Palm (1999) studied the type specimen of Dothiorella pyrenophora, which is the type species of the genus (Sutton 1977), they reported brown and 1-septate conidia of the type that have been associated with Diplodia (Fuckel 1870, Wollenweber 1941). For that reason they regarded Dothiorella as a synonym of Diplodia. We examined the type of Do. pyrenophora (K 54912) and confirm the observations of Crous and Palm (1999). However, Do. pyrenophora has several morphological features that are not consistent with the concept of Diplodia as typified by D. mutila Fr. Thus in Diplodia spp. the conidia initially are hyaline and aseptate but they can become brown and develop one or two eusepta some time after they are formed (Alves et al 2004). This typical form is seen in the anamorphs of B. tsugae A. Funk (a Macrophoma sp.), B. stevensii Shoemaker (Diplodia mutila Fr.), B. quercuum and B. corticola A.J.L. Phillips et al (D. corticola A.J.L. Phillips et al), (Funk 1964, Shoemaker 1964, Alves et al 2004). In contrast conidia of Do. pyrenophora become brown and 1-euseptate while they are still inside the pycnidial cavity and often while attached to the conidiogenous cell (Crous and Palm 1999, present observations). In this respect the anamorphs of the two species described in this paper correlate most closely with Dothiorella as typified by Do. pyrenophora. Furthermore phylogenetically these anamorphs do not belong in Diplodia. For these reasons we consider Dothiorella a suitable genus to accommodate anamorphs of Botryosphaeria species with conidia that become brown and 1-septate soon after they are formed. Because the teleomorphs of the two species described as new in this paper are rare, we believe we are justified in giving names to their anamorphs, which we describe here as Dothiorella sarmentorum and Dothiorella iberica. To clarify the differences between Diplodia and Dothiorella we provide amended descriptions of both genera to include new information that separates them morphologically.
Although Dothiorella appears to be a suitable genus for these anamorphs, species determinations are difficult. In contrast to other Botryosphaeria species, morphological differences are more apparent in the teleomorphs while the anamorphs are morphologically similar and dimensions of the conidia overlap considerably. However by taking into consideration average lengths of conidia and 95% confidence limits, the two species can be separated. Thus, conidia of B. iberica are (17.2–)23.0–23.4(–28.6) x (8.1–) 10.8–11.0(–16) µm and averaged 23.2 x 10.9 µm (n = 400) while in B. sarmentorum they are (17.6–)21.4–21.9(–24.6) x (7.9–)9.7–9.9(–11.6) µm and averaged 21.6 x 9.8 µm (n = 150). The range of dimensions reported by Crous and Palm (1999) for Do. pyrenophora [(12–)25–30(–35) x (7–)11–14(–16) µm] cover the entire range of the isolates we studied but the mean dimensions were 26.5 ± 1.56 x 11.83 ± 1.11 µm, which is larger than both B. iberica and B. sarmentorum.
The genus Dothiorella has been the source of much confusion in the past, and the name has been used in more than one sense. This genus has been used for anamorphs with hyaline aseptate conidia of the type normally associated with Fusicoccum. This confusion presumably started when Petrak (1922) transferred F. aesculi Corda to Dothiorella, citing the species as the conidial state of B. berengeriana (Sutton 1980). In later years Dothiorella has been used for Fusicoccum-like anamorphs with multiloculate conidiomata (Grossenbacher and Duggar 1911, Barr 1987, Rayachhetry 1996). Sivanesan (1984) confused matters further by placing Dothiorella pyrenophora in synonymy with Dothichiza sorbi Lib., which has small, hyaline, aseptate conidia and is the anamorph of Dothiora pyrenophora (Fr.) Fr. However, he was referring to Dothiorella pyrenophora Sacc., 1884, which is a later homonym of Dothiorella pyrenophora Sacc., 1880 (Sutton 1977). The taxonomic history of Dothiorella has been explained by Sutton (1977) and Crous and Palm (1999).
A further matter that developed from this study concerned the relationship between Otthia spiraeae and Diplodia sarmentorum. Booth (1958) connected these two fungi through isolations made from a collection of O. spiraea on Ulmus. This resulted in speculation about the taxonomic position of Otthia and led to suggestions that it should be regarded as a synonym of Botryosphaeria (Laundon 1973, Denman et al 2000). However, Otthia, typified by O. spiraeae, is morphologically distinct from Botryosphaeria. Thus in Otthia, the asci are cylindrical with a thin endotunica while in Botryosphaeria they are clavate with a thick endotunica. Pseudoparaphyses in Otthia are narrow and sinuous, infrequently septate and not constricted at the septa, compared to the broader, frequently septate ones in Botryosphaeria. The specimen that Booth (1958) studied is morphologically distinct from Otthia, and we consider it to be a species of Botryosphaeria, which we describe here as B. sarmentorum. To clarify this aspect we provide a description of O. spiraeae, the lectotype species of Otthia. Finally, we transfer O. quercus to Botryophaeria as B. quercicola nom. nov.
|
TAXONOMIC PART |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONTAXONOMIC PARTLITERATURE CITED |
|---|
= Sphaeropsis Sacc., nom. cons:, Michelia 2:105. 1880.
= Macrophoma (Sacc.) Berl. & Voglino, Atti. Soc. Venet.-Trent. Sci Nat. 10:4. 1886, and Sacc., Syll. Fung. Addit. 1–4:306. 1886.
Phoma Westend. subgen. Macrophoma Sacc., Syll. Fung. 3:65. 1884.
Type species: Diplodia mutila Fr. in Mont., Ann. Sci. Nat. Bot., sér. 2, 1:302. 1834.
Mycelium immersed or superficial, branched, septate, melanized, dark brown. Conidiomata pycnidial, ostiolate, formed in uni- or multiloculate stromata containing up to 20 pycnidial locules each with a prominent ostiole, immersed, becoming erumpent at maturity. Ostiole central, circular, papillate. Paraphyses lacking. Conidiophores (when present) hyaline, simple, occasionally septate, rarely branched, cylindrical, arising from the cells lining the pycnidial cavity. Conidiogenous cells holoblastic, hyaline, cylindrical, determinate or proliferating at the same level giving rise to periclinal thickenings, or proliferating percurrently and forming two or three annellations. Conidia initially hyaline, thick-walled, becoming brown but sometimes the coloration is delayed or never occurs, occasionally becoming 1-euseptate.
Notes. –
The concept of Diplodia has changed over the years and has been regarded as including species with dark brown, 1-septate conidia. However, the genus is typified by D. mutila, which has hyaline, aseptate conidia that can become brown and septate with age. Although Denman et al (2000) regarded Lasiodiplodia as a synonym of Diplodia, and this was accepted by Slippers et al (2004a), there are grounds for regarding them as separate genera. Thus paraphyses and the longitudinal striations seen on the conidium wall are characteristic features of Lasiodiplodia theobromae (Pat.) Griffon & Maubl., the type species of Lasiodiplodia that are not found in any species of Diplodia. Although paraphyses and striate conidia are seen in D. gossypina Cooke, Punithalingam (1976) regarded this species as a synonym of L. theobromae.
Dothiorella Sacc., Michelia 2:5. 1880.
Type species: Dothiorella pyrenophora Sacc., Michelia 2:5. 1880.
Mycelium immersed or superficial, branched, septate, melanized, dark brown. Conidiomata pycnidial, ostiolate, individual or in loose clusters of up to 10 pycnidia, immersed, breaking through the bark when mature. Ostiole circular, central, non-papillate. Paraphyses absent. Conidiophores absent. Conidiogenous cells holoblastic, hyaline, smooth-walled, cylindrical and slightly swollen at the base, determinate or indeterminate and proliferating at the same level to form periclinal thickenings, very rarely proliferating percurrently to produce two or three indistinct annellations, borne directly on the cells lining the pycnidial cavity. Conidia initially hyaline, becoming dark brown and 1-euseptate within the pycnidial cavity and often while still attached to the conidiogenous cell, thick-walled, externally smooth, internally verruculose.
Notes. – This genus differs from Diplodia in the conidia, which in Dothiorella become brown and 1-septate at an early stage of their development and often before they are released from the conidiogenous cell. Conidia in Dothiorella are brown and 1-septate before they are discharged from the pycnidia, while in Diplodia they can become septate and the wall sometimes darkens only some time after discharge from the pycnidium. Furthermore, percurrent proliferation of conidiogenous cells is extremely rare in Dothiorella but common in Diplodia.
Botryosphaeria sarmentorum A.J.L. Phillips, Alves & Luque, sp. nov. FIGS. 2–13
Anamorph: Dothiorella sarmentorum (Fr.) A.J.L. Phillips, Alves & Luque, comb. nov.
= Diplodia sarmentorum (Fr.) Fr., Summ. veg. Scand. (Sweden) 2:417. 1849.
Sphaeria sarmentorum Fr., Syst. mycol. 2:498. 1823.
= Diplodia pruni Fuckel, Jahrb. Nassauischen Vereins Naturk:, 23–24:169. 1870 [1869].
Additional synonyms for Do. sarmentorum according to Wollenweber (1941).
Ascostromata in hospitis inclusa, 350–400 µm diametro, erumpescentia, stromatiformis, atrobrunnea vel nigra, cum ostiolis centralibus, papillatis. Pseudoparaphyses filiformis, septatis. Asci 140–210 x 17–24 µm, stipitati, cylindrici-clavati, quadro- ad octospori, bitunicati cum loculo apicali bene evoluto. Ascosporae irregulariter biseriatae, brunnea, uniseptatae, (21–)22.5–28(–30) x (10–)11–12.5(–14) µm, fusiformis oblongis. Conidiomata in contextu hospitis inclusa, solitaria, stromatiformia, globosa, usque 450 µm diametro. Cellulae conidiogenae holoblasticae, hyalinae, subcylindricae, 7–15 x 3–7 µm, proliferatione percurrenti limitata, ut videtur 2–3 annellationibus paucis, vel inplano eodem periclinaliter incrassate. Conidia brunnea, uniseptata, parietibus crassis, ovoidea, apicibus obtuse rotundato, in fundo truncata, (17.5–)19–23.5(–24.5) x (8–)8.5–10.5(–11.5) µm. Cellulae spermatiogenae 7–10 x 2–3 µm, cylindricae, hyalinae, laevia, holoblasticae, phialidibus typicus periclinaliter spissescentibus. Spermatia hyalina, laevia, unicellulares, 4–5.5 x 2 µm.
Ascomata dark brown to black, globose pseudothecial, 350–400 µm diam, submerged in the substrate, partially erumpent at maturity, ostiolate; ostiole circular, central, papillate; wall 50–75 µm thick, composed of dark brown thick-walled textura angularis, cells 10–17 x 6–9 µm, lined with thinner-walled, hyaline, textura angularis. Pseudoparaphyses thin-walled, hyaline, frequently septate, often constricted at the septa, 3–4 µm wide. Asci 140–210 x 17–24 µm, stipitate, arising from the base of the ascoma, cylindric-clavate, bitunicate, endotunica thick-walled, with a well developed apical chamber, 4–6(–8)-spored, obliquely uniseriate or irregularly biseriate. Ascospores (21.2–)24.4–25.5(–30.5) x (9.9–)11.8–12.3(–14.0) µm, mean ± SD of 50 ascospores = 25.0 ± 2.0 x 12.1 ± 0.9 µm, oblong to ovate, widest in the middle part, straight, (0–)1 septate, slightly constricted at the septum, dark brown, moderately thick-walled, surface smooth, finely verruculose on the inner surface. Conidiomata solitary, stromatic, globose, up to 450 µm wide, wall 5–8 cell layers thick, composed of dark brown thick-walled textura angularis, becoming thin-walled and hyaline toward the inner region. Conidiophores reduced to conidiogenous cells. Conidiogenous cells lining the pycnidial cavity, holoblastic, hyaline, subcylindrical, 7–15 x 3–7 µm, proliferating at the same level giving rise to periclinal thickenings or rarely proliferating percurrently to form one or two close, indistinct annellations. Conidia (17.6–)21.4–21.9(–24.6) x (7.9–)9.7–9.9(–11.6) µm, mean ± SD of 150 conidia = 21.6 ± 1.5 x 9.8 ± 0.9 µm, L/W ratio = 2.2, brown walled, 1-septate, slightly constricted at the septum, ovoid with a broadly rounded apex and truncate base. Spermatiogenous cells discrete or integrated, hyaline, smooth, cylindrical, holoblastic or proliferating via phialides with periclinal thickenings, 7–10 x 2–3 µm. Spermatia hyaline, smooth, aseptate, rod-shaped with rounded ends, 4–5.5 x 2 µm.
Cardinal temperatures for growth. min 5 C, opt 20–25 C, max below 35 C.
Known hosts. – Plurivorous including Malus, Menispermum, Prunus, Pyrus, Ulmus.
Known geographical range. – England, The Netherlands, Norway, Sweden, possibly worldwide.
HOLOTYPE of B. sarmentorum: ENGLAND. WARWICKSHIRE. On Ulmus sp., Aug. 1956, E. A. Ellis (IMI 63581b, as Otthia spiraeae).
Specimens examined. – Sphaeria sarmentorum; SWEDEN: Lund, Botanical Garden. On Menispermum canadense, 1818, E. M. Fries Scleromyc. Suec. 18 (HOLOTYPE for the anamorph UPS-FRIES; ISOTYPE for the anamorph K(M) 104852). Diplodia pruni; GERMANY. On Prunus armeniaca, collection date unknown, K. W. G. L. Fuckel, Fung. Rhen. No. 1710 (HOLOTYPE in G).
Cultures examined. – Dothiorella sarmentorum; LOCATION UNKNOWN. On Prunus armeniaca, Jan 1933, R. M. Nattrass (CBS 165.33, as Diplodia pruni). NORWAY: On Pyrus communis, Aug 1941, H. W. Wollenweber (CBS 120.41, as Diplodia sarmentorum). ENGLAND. WARWICKSHIRE. On Ulmus sp., Aug 1956, E. A. Ellis (IMI 63581b, as Diplodia sarmentorum, culture ex type of B. sarmentorum). THE NETHERLANDS: Delft. On Malus pumila, Apr 2003, A. J. L. Phillips (CBS 115038).
Notes. –
In proposing 145 species as synonyms of Do. sarmentorum, Wollenweber (1941) reported a wide range of dimensions for the conidia, namely, (15–) 20–24(–35) x (7–)7.4–11.5(–15) µm. As shown in the present study, some species in Dothiorella are separated by minor differences in conidium dimensions. It therefore is possible that some of Wollenweber’s synonyms are in fact distinct species.
Botryosphaeria iberica A.J.L. Phillips, Luque & Alves, sp. nov. FIGS. 14–27
Anamorph: Dothiorella iberica A.J.L. Phillips, Luque & Alves, sp. nov.
Ascostromata in hospitis inclusa, usque ad 350 µm diametro, erumpescentia, stromatiformis, atrobrunnea vel nigra, cum ostiolis centralibus, papillatis. Pseudoparaphyses filiformis, septatis. Asci 100–125 x 18–25 µm, stipitati, cylindrici-clavati, quadro- ad octospori, bitunicati cum loculo apicali bene evoluto. Ascosporae irregulariter biseriatae, brunneae, uniseptatae, (18.0–)19.5–25.5(–29) x (8.5–)9.0–11.5(–12.5) µm, fusiformis oblongis vel subclavatis, apicibus obtusis, in fundo subacutis. Conidiomata in contextu hospitis inclusa, solitaria, stromatiformia, globosa, usque 450 µm diametro. Cellulae conidiogenae holoblasticae, hyalinae, subcylindricae, 8–15 x 3–5(–6.5) µm, proliferatione percurrenti limitata, ut videtur 2–3 annellationibus paucis, vel inplano eodem periclinaliter incrassate. Conidia brunnea, uniseptata, parietibus crassis, ovoidea, apicibus obtuse rotundato, in fundo truncata, (21.5–)23–26(–27.5) x (8.5–) 9–12.5(–13.5) µm. Spermatia non visa.
Ascomata dark brown to black, globose pseudothecial, up to 350 µm diam, submerged in the substrate, partly erumpent at maturity, ostiole circular, central, papillate; wall up to 50 µm thick, composed of dark brown thick-walled textura angularis, cells 8–17 x 6–10 µm and lined with thinner-walled, hyaline, textura angularis. Pseudoparaphyses thin walled, hyaline, frequently septate, slightly constricted at the septum, 2.5–3.5(–4) µm wide. Asci 100–125 x 18–25 µm, stipitate, arising from the base of the ascoma, clavate, thick-walled, bitunicate with a well-developed apical chamber, stipitate, (4–)8-spored, irregularly biseriate. Ascospores (17.8–)22.5–23.7(–29.2) x (8.7–) 10.0–10.4(–12.3) µm, mean ± SD of 50 ascospores = 23.1 ± 2.1 x 10.2 ± 0.8 µm, oblong, ovate to subclavate, (0–)1 septate, slightly constricted at the septum, dark brown, moderately thick-walled, finely verruculose on the inner surface, straight or inequilateral, widest in the lower one-third to middle of the apical cell, basal cell tapering toward the rounded end. Conidiomata solitary, stromatic, globose, up to 450 µm wide, thick walled, composed of dark brown thick-walled textura angularis, becoming thin-walled and hyaline toward the inner region. Conidiophores reduced to conidiogenous cells. Conidiogenous cells lining the cavity, holoblastic, hyaline, subcylindrical, 8–15 x 3–5(–6.5) µm, proliferating at the same level giving rise to periclinal thickenings or rarely proliferating percurrently forming one or two indistinct annellations. Conidia (17.2–)23.0–23.4(–28.6) x (8.1–)10.8–11.0(–16.0) µm, mean ± SD of 400 conidia 23.2 ± 1.9 x 10.9 ± 1.2 µm, L/W ratio = 2.2, brown walled, 1-septate, slightly constricted at the septum, ovoid with a broadly rounded apex and truncate base. Spermatia not seen.
Cardinal temperatures for growth. min 5 C, opt 20–25 C, max below 35 C.
Known hosts. – Malus, Quercus.
Known geographical range. – Italy, Spain.
Specimens examined. – SPAIN. ARAGON: Tarazona. On dead twigs of Quercus ilex, 18 Dec 2002, J. Luque, (HOLOTYPE of B. iberica, LISE 94944, culture ex type CBS 115041). SPAIN. ARAGON: Monzón. On dead twigs of Quercus ilex, 24 Sep 1999, N. Ibarra (HOLOTYPE of Do. iberica, LISE 94942, culture ex type CBS 115035).
Cultures examined. –
(TABLE I.)
Notes. – This species is similar to B. sarmentorum but can be distinguished on characteristics of the asci, ascospores and conidia. Thus in B. iberica the asci are shorter and more clavate, the ascospores characteristically taper toward the base, and on average the conidia are slightly longer.
Botryosphaeria quercicola A.J.L. Phillips, nom. nov. FIGS. 28–36
|
Otthia quercus Fuckel, Jahrb. Nassauischen Vereins Naturk., 23–24:170. 1870 [1869]. ?Anamorph: Diplodia quercus Fuckel, Jahrb. Nassauischen Vereins Naturk., 23–24:170. 1870 [1869].
Ascomata dark brown to black, globose pseudothecia, multiloculate, individual locules 250–300 µm diam, immersed in the host and erumpent at maturity, ostiolate, ostioles central, circular, papillate; wall up to 100 µm thick, composed of up to 12 layers of thick-walled textura angularis, cells 17–34 x 10–18 µm, inner layers hyaline, thin-walled and flattened. Pseudoparaphyses 3.6–4.8 µm wide, thin-walled, hyaline, frequently septate, constricted at septum. Asci 110–165 x 27.5–38 µm, arising from base of ascoma, cylindrical to clavate, thick-walled, bitunicate with a well developed apical chamber, stipitate, 8-spored, irregularly biseriate. Ascospores brown, (28.3–)30.7–33.9(–35.1) x (10.4–)11.8–13.9(–15.5) µm, mean ± SD of 30 ascospores = 32.3 ± 2.5 x 12.9 ± 1.6 µm, 1–2-septate at maturity, constricted at septum, thin-walled, wall internally roughened, oval to broadly fusiform, both ends rounded, widest in the middle or upper third. Conidiomata eustromatic, uni- or multiloculate, ostiolate, ostiole central, papillate. Conidiogenous cells cylindrical, discrete, indeterminate and proliferating percurrently to produce 2–3 annellations or proliferating at the same level to form periclinal thickenings. Conidia (24–)28.8–30.8(–38) x (11–)15.9–17.1(–21.2) µm, mean ± SD of 50 conidia = 29.8 ± 3.6 x 16.5 ± 2.1 µm, L/W ratio = 2.1, hyaline, aseptate, moderately thick walled, wall smooth, eguttulate, oval, both ends rounded. Spermatiogenous cells discrete or integrated, hyaline, smooth, cylindrical, holoblastic or proliferating via phialides with periclinal thickenings, 10–18 x 1.5–4 µm. Spermatia hyaline, smooth, aseptate, rod-shaped with rounded ends, 4.5–9 x 1.5–2 µm.
Known hosts. – Quercus species.
Known geographical range. – Germany.
Specimen examined. – GERMANY: Hessen. On Quercus, date unknown, K. W. G. L. Fuckel, Fungi Rhenani 534 (HOLOTYPE in G).
Notes. –
A new name is proposed rather than a new combination since B. quercus could be confused with B. quercuum. The description of D. quercus was taken from Fungi Rhenani 534. Although Fuckel regarded this as the anamorph of O. quercus, the connection between the two has not been demonstrated definitively through culture of ascospores. Diplodia quercus is similar to Diplodia corticola except that conidia of the latter are wider (Alves et al 2004).
Otthia spiraeae (Fuckel) Fuckel, Jahrb. Nassauischen Vereins Naturk., 23–24:170. 1870 [1869]. FIGS. 37–41
Cucurbitaria spiraeae Fuckel, Fungi Rhenani Exs. No. 975 (1863).
Anamorph: Not known.
Ascomata stromatic, dark brown to black, subcortical, necks erumpent at maturity; wall 60–70 µm wide, composed of 10–12 layers of thick-walled textura angularis, cells 7–12 x 5–10 µm. Pseudoparaphyses 1.5–2.5 µm wide, thin walled, hyaline, sinuous, unbranched, infrequently septate, not constricted at septum. Asci 210–230 x 19–21 µm, cylindrical, bitunicate, containing eight obliquely uniseriate ascospores. Ascospores (22.2–)24.6–25.3(–29.5) x (10.5–) 11.5–11.9(–13.9) µm, mean ± SD of 30 ascospores = 25.0 ± 1.2 x 11.7 ± 0.7 µm, dark brown, 1-septate, slightly constricted at the septum, oval, both ends rounded, moderately thick-walled.
Specimens examined. – Cucurbitaria spiraeae; GERMANY. On dry twigs of Spiraea opulifolia, date unknown, K. W. G. L. Fuckel, Fung. Rhen. No. 975 (HOLOTYPE K 104853).
|
ACKNOWLEDGMENTS |
|---|
|
FOOTNOTES |
|---|
1 Corresponding author. E-mail: alp{at}mail.fct.unl.pt
|
LITERATURE CITED |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONTAXONOMIC PARTLITERATURE CITED |
|---|
Anonymous. 1968. Plant Pathologists Pocketbook. Kew, England: CMI. 267 p.
Barr ME. 1972. Preliminary studies on the Dothideales in temperate North America. Contr Univ Michigan Herb 9:523–638.
———. 1987. Prodromus to Class Loculoascomycetes. Amherst, Massachusetts: Publ. by the author. 168 p.
———. 1989. The genus Dothidotthia (Botryosphaeriaceae) in North America. Mycotaxon 34:517–526.
Booth C. 1958. Studies of pyrenomycetes III. Otthia spiraeae (Fuckel) Fuckel, syn. Diplodia sarmentorum (Fr.) Fr. Trans Br Mycol Soc 41:335–340.
Carbone I, Kohn LM. 1999. A method for designing primer sets for speciation studies in filamentous ascomycetes. Mycologia 91:553–556.[CrossRef]
Cesati V, De Notaris G. 1863. Schema di classificazione degle sferiacei italici aschigeri piu’ o meno appartenenti al genere Sphaeria nell’antico significato attribuito gli de Persoon. Comm Soc Crittogam Ita. 1 , 4:177–240.
Crous PW, Palm ME. 1999. Reassessment of the anamorph genera Botryodiplodia, Dothiorella and Fusicoccum. Sydowia 51:161–175.
Denman S, Crous PW, Taylor JE, Kang J-C, Pascoe I, Wingfield MJ. 2000. An overview of the taxonomic history of Botryosphaeria, and a re-evaluation of its anamorphs based on morphology and ITS rDNA phylogeny. Stud Mycol 45:129–140.
———, ———, Groenewald JZ, Slippers B, Wingfield BD, Wingfield MJ. 2003. Circumscription of Botryosphaeria species associated with Proteaceae based on morphology and DNA sequence data. Mycologia
95:294–307.
Eriksson OE. 1981. The families of bitunicate Ascomycetes. Opera Botanica 60:1–220.
Fuckel L. 1870. Symbolae mycologicae. Jb. Nassau. Ver. Naturk. 23–24:1–459.
Funk A. 1964. Botryosphaeria tsugae n. sp. causing dieback of western hemlock in British Columbia. Can J Bot 42: 769–775.
Grossenbacher JG, Duggar BM. 1911. A contribution to the life history, parasitism, and biology of Botryosphaeria ribis. New York Agric Exp Stn Geneva Tech Bull 18: 114–188.
Hillis DM, Bull JJ. 1993. An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Syst Biol 42:182–192.[CrossRef]
Jacobs KA, Rehner SA. 1998. Comparison of cultural and morphological characters and ITS sequences in anamorphs of Botryosphaeria and related taxa. Mycologia 90:601–610.[CrossRef]
Laundon GF. 1973. Botryosphaeria obtusa, B. stevensii and Otthia spiraeae in New Zealand. Trans Brit Mycol Soc 6:369–374.
Morgan-Jones G, White JF Jr. 1987. Notes on Coelomycetes. II. Concerning the Fusicoccum anamorph of Botryosphaeria ribis. Mycotaxon 30:117–125.
Pennycook SR, Samuels GJ. 1985. Botryosphaeria and Fusicoccum species associated with ripe fruit rot of Actinidia deliciosa (Kiwifruit) in New Zealand. Mycotaxon 24: 445–458.
Petrak F. 1922. Beiträge zur Kenntnis der Pilzflora der südlichen Alpenländer und Norditaliens. Ann Mycol 20: 126–159.
Phillips AJL, Fonseca F, Povoa V, Castilho R, Nolasco G. 2002. A reassessment of the anamorphic fungus Fusicoccum luteum and description of its teleomorph Botryosphaeria lutea sp. nov. Sydowia 54:59–77.
Punithalingam E. 1976. Botryodiplodia theobromae. CMI descriptions of pathogenic fungi and bacteria, No. 519. Kew, Surrey, England: Commonwealth Mycological Institute. 2 p.
Rayachhetry MB, Blakeslee GM, Webb RS, Kimbrough JW. 1996. Characteristics of the Fusicoccum anamorph of Boryosphaeria ribis, a potential biological control agent for Melaleuca quinquenervia in South Florida. Mycologia 88:239–248.[CrossRef]
Saccardo PA. 1877. Fungi veneti novi vel critici vel Mycologiae Venetae addendi. Michelia 1:1–72.
Samuels GJ, Singh B. 1986. Botryosphaeria xanthocephala cause of stem canker in pigeon pea. Trans Br Mycol Soc 86:295–299.
Shoemaker RA. 1964. Conidial states of some Botryosphaeria species on Vitis and Quercus. Can J Bot 42:1297–1303.
Sivanesan A. 1984. The bitunicate ascomycetes and their anamorphs. Vaduz, Liechtenstein: J Cramer. 701 p.
Slippers B, Crous PW, Denman S, Coutinho TA, Wingfield BD, Wingfield MJ. 2004a. Combined multiple gene genealogies and phenotypic characters differentiate several species previously identified as Botryosphaeria dothidea. Mycologia
96:83–101.
———, Fourie G, Crous PW, Coutinho TA, Wingfield BD, Wingfield MJ. 2004b. Multiple gene sequences delimit Botryosphaeria australis sp. nov. from B. lutea. Mycologia
96:1030–1041.
Smith H, Crous PW, Wingfield MJ, Coutinho TA, Wingfield BD. 2001. Botryosphaeria eucalyptorum sp. nov., a new species in the B. dothidea-complex on Eucalyptus in South Africa. Mycologia 93:277–285.[CrossRef]
Stevens NE. 1936. Two species of Physalospora in England. Mycologia 28:330–336.[CrossRef]
Sutton BC. 1977. Coelomycetes VI. Nomenclature of generic names proposed for Coelomycetes. Mycol Pap 141: 1–253.
———. 1980. The Coelomycetes. Kew, England: CMI. 696 p.
Swofford DL. 2003. PAUP*. Phylogenetic analysis using parsimony (*and other methods) version 4. Sunderland, Massachusetts: Sinauer Associates.
Theissen F, Sydow H. 1915. Die Dothideales. Ann Mycol 13: 149–746.
van Niekerk JM, Crous PW, Groenewald JZ, Fourie PH, Haleen F. 2004. DNA phylogeny morphology and pathogenicity of Botryosphaeria species occurring on grapevines. Mycologia
96:781–798.
von Arx JA, Müller E. 1954. Die Gattungen der amerosporen Pyrenomyceten. Beitr Kryptfl Schweiz 11(1):1–434.
von Höhnel F. 1909. Fragmente zur Mykologie. Sitzungsb Kaiserl Akad Wiss, Math-Naturwiss Kl, Abt. 1 , 118:813–904.
White TJ, Bruns T, Lee S, Taylor J. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ, eds. PCR protocols: a guide to methods and applications. San Diego, California: Academic Press. p 315–322.
Wollenweber HW. 1941. Diplodia sarmentorum Fries und ihre Vernreitung. Zentralbl Bakteriol Parasitendk 103: 347–357.
Zhou S, Stanosz GR. 2001. Relationships among Botryosphaeria species and associated anamorphic fungi inferred from the analyses of ITS and 5.8s rDNA sequences. Mycologia 93:516–527.[CrossRef]
This article has been cited by other articles:
 |
|
 |
|
  E. I. Rojas, E. A. Herre, L. C. Mejia, A. E. Arnold, P. Chaverri, and G. J. Samuels Endomelanconiopsis, a new anamorph genus in the Botryosphaeriaceae Mycologia, September 1, 2008; 100(5): 760 - 775. [Abstract] [Full Text] [PDF]
|
|
|
|
|
|
T. I. Burgess, P. A. Barber, S. Mohali, G. Pegg, W. de Beer, and M. J. Wingfield Three new Lasiodiplodia spp. from the tropics, recognized based on DNA sequence comparisons and morphology. Mycologia, May 1, 2006; 98(3): 423 - 435. [Abstract] [Full Text] [PDF]
|
|
|
|
 |
|
 P. W. Crous, B. Slippers, M. J. Wingfield, J. Rheeder, W. F.O. Marasas, A. J.L. Philips, A. Alves, T. Burgess, P. Barber, and J. Z. Groenewald Phylogenetic lineages in the Botryosphaeriaceae. Stud Mycol, January 1, 2006; 55: 235 - 253. [Abstract] [Full Text] [PDF]
|
|
|
|
|
|
J. Luque, S. Martos, and A. J.L. Phillips Botryosphaeria viticola sp. nov. on grapevines: a new species with a Dothiorella anamorph. Mycologia, September 1, 2005; 97(5): 1111 - 1121. [Abstract] [Full Text] [PDF]
|
|
|
|
|
|
D. F. Farr, M. Elliott, A. Y. Rossman, and R. L. Edmonds Fusicoccum arbuti sp. nov. causing cankers on Pacific madrone in western North America with notes on Fusicoccum dimidiatum, the correct name for Scytalidium dimidiatum and Nattrassia mangiferae Mycologia, May 1, 2005; 97(3): 730 - 741. [Abstract] [Full Text] [PDF]
|
|
| ||||||||
