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Departamento de Fitopatologia, Universidade de Brasília, 70910-900 Brasília, D.F., Brasil
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ABSTRACT |
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New hyphomycetes are described in association with leaves of native plants of the family Vochysiaceae, as part of studies of cerrado fungi. Six new species are described belonging to genera Alternaria (A. qualeae sp. nov.), Janetia (J. salvertiae sp. nov.), Passalora (P. qualeae sp. nov.) and Periconiella (P. longispora sp. nov., P. qualeae-grandiflorae sp. nov. and P. campo-grandensis sp. nov.). A key to the species of Periconiella on Qualea is provided.
Key words: tropical fungi
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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, 1993). The main genus in the family is Qualea Mart., however species of Salvertia St. Hil. and Vochysia Aublet also are found frequently (Stafleu 1948, 1953; Joly 1993; Paula and Alves 1997). The widespread presence of the family in the cerrado is connected to its ability to tolerate and accumulate aluminum, which is present in toxic levels in the soils of the region (Goodland 1971, Haridasan 1982, Araújo 1984).
A diverse and large population of plant-associated fungi is present in the cerrado (Dianese et al 1997, 2001; Dianese 2000). In the Vochysiaceae, 32 fungal species have been recorded, among them are 25 foliicolous Ascomycota. Ten species have been found on Qualea, 12 on Vochysia and two on Salvertia species (Mendes et al 1998). Five of the recently described fungal species on Qualea were three Uncinula species (Dianese and Dianese 1995), a rust, Aplopsora hennenii Dianese & Santos (Dianese and Santos 1995) and the coelomycete Harknessia salvertiana Furlanetto & Dianese on Salvertia convallariodora (Furlanetto and Dianese 1998).
The aim of the present work was to study the fungi associated with about 600 specimens of the Vochysisaceae deposited in the Mycological Reference Collection of the Herbarium of the University of Brasília. New taxa on Qualea, Vochysia and Salvertia species are described herein.
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MATERIALS AND METHODS |
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Slides containing squash preparations of fungal fruiting bodies or sections made with a freezing microtome were used for morphological studies and microphotography. In most cases, the samples were stained with lacto-glycerol-cotton blue or glycerol-KOH-phloxine B and the slides sealed with nail polish.
Pieces of leaves with one or more lesions showing representative samples of fruiting bodies were used for scanning electron microscopy (SEM) observation after being fixed in sodium caccodylate buffer, pH 7·4, 0·1M, containing glutaraldehyde 2%, for at least 24 h. The samples were dehydrated in an aqueous series with increasing acetone concentrations from 15, 30, 50, 75 up to 100%, for 15 min in each concentration. Leaf pieces were dried at the critical point before being covered by a thin layer of gold with a sputter coater for 2 min (Souza 1998). Samples were observed with a scanning electron microscope (Jeol, model JSM 840-A E).
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TAXONOMY |
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Coloniae amphigenae, effusae, velutinae, pallido brunneae vel atro-brunneae. Mycelium brunneum, superficiale, ex parte immersum. Hyphae 4–5 µm diam, brunneae, septatae, ramosae. Stromata subcuticulares vel erumpentia, textura angularis. Conidiophora adaxialia 42–132 (75) x 5–8 (5) µm, fasciculata, mononematica, erecta, sympodialia, porosa, 3, 4, vel 5-plo longia quam conidiophora abaxialia. Conidiophora abaxialia 8–25 (17) x 5–8 (5) µm, non-fasciculata, solitaria, mononematica, sympodialia, porosa, velut rami laterales hyphae superficialium vel apices hyphae formata. Cellulae conidiogenae integratae, sympodiales, enteroblasticae, polytreticae, cum cicatricibus porosis. Conidia 20–83 (43) x 9–16 (12) µm, muriformes, atro brunnea, rostrata. Rostrum pallido brunneum, saepe dimidium conidia longior.
Colonies amphigenous, effuse, velutinous, pale to dark brown. Mycelium brown, superficial, partially immersed. Hyphae 4–5 µm diam, septate, branched. Stromata subcuticular becoming erumpent, textura angularis (Figs. 1, 2, 4, 5). Adaxial conidiophores 42–132 (75) x 5–8 (5) µm, fasciculate, mononematous, occasionally branched, erect, sympodially proliferating, 3–5 times longer than the abaxial conidiophores (Figs. 1–3, 6). Abaxial conidiophores 8–25 (17) x 5–8 (5) µm, single, mononematous, sympodially proliferating, originating laterally or apically on superficial hyphae (Figs. 7, 8). Conidiogenous cells integrated, sympodial, enteroblastic, polytretic, with porous scars (Figs. 8, 9). Conidia 20–83 (43) x 9–16 (12) µm, muriform, dark brown, rostrate. Beak pale brown, mostly equal or longer than half of the conidial length (Figs. 7, 8, 10).
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Alternaria Nees ex Fries contains about 270 species with muriform, pale brown to brown, obclavate, mostly rostrate, catenulate or non-catenulate conidia. The conidiogenesis is typically enteroblastic polytretic (Ellis 1971, 1976; Kirk et al 2001).
Simmons (1981, 1982a, 1982b, 1990, 1993a, 1993b, 1994a, 1994b, 1997, 1999) and Simmons and Roberts (1993) have been studying the genus Alternaria, describing new species and revising existing taxa, using host or plant family as one of the primary taxonomic characters. The 12 families in the order Rutales include Rutaceae and Vochysiaceae. However, only in the genus Citrus of family Rutaceae was Alternaria species previously recorded (Viégas 1961, Silva and Minter 1995, Ellis 1976) and recently revised (Simmons 1999). These pathogenic species associated with citrus, all with catenate conidia, were accepted or described by Simmons (1999): A. alternata (Fr.) Keissler, A. limoniasperae Simmons, A. citrimacularis Simmons, A. tangelonis Simmons, A. citriarbusti Simmons, A. turkisafria Simmons, A. toxicogenica Simmons, A. colombiana Simmons, A. perangusta Simmons, A. interrupta Simmons and A. dumosa Simmons. Although Simmons (1999) based his studies mostly on cultural characters, all mentioned species clearly were different from A. qualeae by forming long chains of conidia. Furthermore, conidiophores grouped on adaxially located stromata versus sporulation limited to superficial mycelium on the abaxial face, plus noncatenate or very short (generally two conidia) chains of rostrate conidia, are distinctive characters present in A. qualeae. Finally, the morphological characteristics described linked with the fact that this is the first record of an Alternaria species on vochysiaceous plants lead to the conclusion that the specimen studied here must be accommodated as a new taxon, A. qualeae.
Janetia salvertiae Dornelo-Silva & Dianese, sp. nov. Figs. 11–16
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Coloniae effusae, atro brunneae vel nigrae, velutinae, hypophyllae. Mycelium superficiale, atro brunneum, trichomati insidenti. Hyphae 4 µm diam, septatae, ramosae, cum parietibus laevibus. Conidiophora 10–35 (15) x 4–5 (5) µm, micronematica, mononematica, erecta, sporodochialia. Sporodochium trichomati insidenti. Cellulae conidiogenae 8–15 x 4–5 µm, integratae, saepe intercalares, polyblasticae, 1–2 denticulatae. Conidia 15–30 (20) x 3–5 (4) µm, solitaria, sicca, cylindrica vel clavata, pallido brunnea, 1–6 septatae, cum basibus truncatis reliquiis parietum involucratis.
Colonies effuse, dark brown to blackish, velutinous, hypophyllous. Mycelium superficial, dark brown, on trichomes (Figs. 11–15). Hyphae 4 µm diam, septate, branched, with smooth walls. Conidiophores 10–35 (15) x 4–5 (5) µm, micronematous, erect, sporodochial (Figs. 13–15). Sporodochia on trichomes (Figs. 13–15). Conidiogenous cells 8–15 x 4–5 µm, integrated, mostly intercalary, polyblastic, 1–2 denticulate (Fig. 14). Conidia 15–30 (20) x 3–5 (4) µm, single, dry, cylindrical to clavate, light brown, 1–6-septate, with truncate base containing remnants of the conidiogenous cell wall (rhexolitic secession) (Figs. 14–16).
Specimens examined. BRAZIL. GO: Cristalina, Fazenda Vereda do Gato, on living leaves of Salvertia convallariaeodora, 23 May 1995, M Sanchez n. 829 (HOLOTYPUS: UB col. micol. 8578). BRAZIL. GO: Cristalina, Fazenda Vereda do Gato, on Salvertia convallariaeodora, 23 May 1995, M Sanchez n. 830 (UB col. micol. 8579). BRAZIL. GO: Cristalina, Fazenda Vereda do Gato, on leaves of Vochysia sp. 23 May 1995, JC Dianese n. 2306 (UB col. micol. 8524).
In Janetia M. B. Ellis 17 species have been described, characterized by conidiogenous cells which are micronematous, denticulate, intercalary, integrated, and polyblastic (Ellis 1976, Hughes and Cavalcanti 1983, Goh and Hyde 1996, Calduch et al 2002). Janetia euphorbiae M. B. Ellis on stems of Euphorbia tirucallis L. and J. mangiferae Hughes and Cavalcanti on petioles and branches of Mangifera indica L. are the species that show some similarity to J. salvertiae, mainly due to the formation of 1–2-denticulate conidiogenous cells and 3–6-septate conidia in the case of J. euphorbiae. However, J. salvertiae has conidiophores with 1–2 integrated conidiogenous cells and produces shorter and narrower conidia than those of J. euphorbiae (18–36 [28] x 6–8 µm). Janetia mangiferae, with wider but shorter conidia (8.5–23 x 4.3–6 µm), conidiogenous cells with up to 6 denticles, and close association with an ascomycete (Stomiopeltis Theiss) also is different from the new species. The formation of sporodochia on host trichomes is another unique character of the new species J. salvertiae. The evidence discussed shows that the fungus on S. convallariaeodora constitutes a second record of the genus Janetia in Brazil and is now accommodated in a new species, J. salvertiae.
Passalora qualeae Dornelo-Silva & Dianese, sp. nov. Figs. 17–25
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Coloniae hypophyllae, effusae, velutinae, atro brunneae, irregulares, omnino pagina folii dispositae. Mycelium immersum. Hyphae septatae, ramosae, brunneae. Stromata 35–104 µm diam, atro brunnea, subepidermalia, erumpentia, textura angularis. Conidiophora 92–242 (170) x 5–9 (7) µm, brunnea, macronematica, mononematica, fasciculata, caespitosa, laevia, flexuosa vel erecta, non ramosae. Cellulae conidiogenae 15–25(19) x 5–9 (7) µm, integratae, saepe terminales, aliquando intercalares, polyblasticae, sympodiales, cum cicatricibus fuscatis. Conidia 35–95 (75) x 5–9 (7) µm, 1–3 septata, pallido brunnea, obclavata, solitaria, verruculosa, guttulata, sicca, acropleurogena.
Colonies hypophyllous, effuse, velutinous, dark brown, irregular, sometimes covering the entire leaf blade. Mycelium immersed. Hyphae septate, branched, brown. Stromata 35–104 µm diam, dark brown, subepidermal, erumpent, textura angularis (Figs. 17, 18). Conidiophores 92–242 (170) x 5–9 (7) µm, brown, macronematous, mononematous, fasciculate, caespitose, smooth, flexuous or straight, single (Figs. 17–19, 22, 23). Conidiogenous cells 15–25 (19) x 5–9 (7) µm, integrated, terminal becoming intercalary, polyblastic, sympodial, with darkened scars (Figs. 19, 20, 23, 24). Conidia 35–95 (75) x 5–9 (7) µm, 1–3 septate, light brown, obclavate, solitary, verruculose, guttulate, dry, acropleurogenous (Figs. 21, 25).
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In Brazil, Passalora species have been found in more than 20 host species on plants belonging in families Palmae, Myrtaceae and Gramineae (Mendes et al 1998, Crous et al 1997, Crous and Câmara 1998). The specimen studied conforms well with the concept of Passalora in Braun (1995, 1998), which accommodates cercosporoid fungi with internal mycelium, with stomatal or subepidermal stromata bearing fascicles of pale brown to brown conidiophores containing conidiogenous cells with flat scars, which are darkened and which support single multiseptate pigmented conidia. Following those criteria, Crous et al (1997) and Crous and Câmara (1998) moved eight Cercospora species to Passalora, following Braun (1995, 1998). We have adoped this broader concept instead of that of Deighton (1967), Ellis (1971, 1976) and Medeiros and Dianese (1994) and have concluded that the fungus is a Passalora species. Comparison of P. qualeae with the currently known species of the genus (Deighton 1967; Ellis 1971, 1976; Medeiros and Dianese 1994; Crous et al 1997; Crous and Câmara 1998) indicate that the fungus is a new species and the first described species on a vochysiaceous host.
New Periconiella species
Among the hyphomycetes on Qualea grandiflora three new Periconiella species were found and are described here.
Periconiella longispora Dornelo-Silva & Dianese, sp. nov. Figs. 26–32
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Coloniae hypophyllae, cinerascentes, velutinae, effusae, intercostales. Mycelium superficiale, ex parte immersum. Hyphae septatae, ramosae, hyalinae vel pallido brunneae, laeviae. Conidiophora 300–770 (514) x 7–10 µm, macronemata, erecta, raro curvata, brunnea vel atro brunnea, septata, laevia, cum cellulis basalibus podiformibus, cum apice et subapice ramosi, rami saepe primari, rami secundari raro. Rami primarii 60–137 (94) x 7–9 µm, ramos secundarios vel cellulas conidiogenas ferentes. Rami secundari cylindrici, breves. Cellulae conidiogenae ampuliformes vel lageniformes, polyblasticae, integratae, sympodiales, cicatricatae. Conidia 37–95 (53) x 5–8 (5) µm, acropleurogena, solitaria, cylindrica, pallido brunnea vel brunnea, 3–5 septata, laevia, cum basibus truncatis.
Colonies hypophyllous, grayish, velutinous, effuse, growing among leaf ribs. Mycelium superficial, partially immersed. Hyphae septate, branched, hyaline to light brown, smooth. Conidiophores 300–770 (514) x 7–10 µm, macronematous, straight, seldom curved, brown to dark brown, septate, smooth, with a defined foot cell, apex and subapex branched, with primary branches in large numbers, secondary branches rare (Figs. 26, 27). Primary branches 60–137 (94) x 7–9 µm, bearing secondary branches or conidiogenous cells (Figs. 28, 30). Secondary branches cylindrical, short. Conidiogenous cells ampulliform or lageniform, polyblastic, integrated, sympodial, cicatrized (Figs. 29, 30). Conidia 37–95 (53) x 5–8 (5) µm, acropleurigenous, single, cylindrical, light brown to brown, 3–5-septate, smooth, with truncate base (Figs. 31, 32).
Specimen examined. BRAZIL. DF: Planaltina, Estação Ecologica de Águas Emendadas, on living leaves of Qualea grandiflora, 17 Sep 1995, M Sanchez n. 1248 (HOLOTYPUS: UB col. micol. 10042).
The main taxonomic features applied for separation of species within the genus Periconiella are conidiophore size, conidial dimensions, septation and ornamentation, septation and dimension of primary, secondary or tertiary apical branches of the conidiophores (Ellis 1967, 1971, 1976). Most of the 48 Periconiella species known show 0–3 septate conidia (Ellis 1967, 1971, 1976; Priest 1991; McKenzie 1996). Among the species showing conidia with three or more septa that could be confused with P. longispora, there are only two (Ellis 1967)—P. santoloides Ellis and P. rapaneae Ellis. Periconiella longispora differs from P. santoloides by verrucose shorter conidia possessing a maximum of four conidial septa, as does P. rapaneae. The latter species further differs from the new species by longer conidiophore primary branches (170 x 5–8 µm) and the presence of tertiary branches. Thus the specimen on Qualea grandiflora is placed in a new species, P. longispora.
Periconiella qualeae-grandiflorae Dornelo-Silva & Dianese, sp. nov. Figs. 33–36
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Coloniae hypophyllae, cinerascentes, velutinae, intercostales. Mycelium superficiale ex parte immersum. Hyphae septatae, ramosae, hyalinae vel pallido brunneae, laeviae. Conidiophora 250–520 (320) x 7–11 (9) µm, macronemata, mononemata, erecta, raro curvata, brunnea vel atro brunnea, septata, laevia, cum ramis primariis treintibus apicalibus stipitum locatis. Rami primarii 23–69 (43) x 10 µm, 2–6-cellulis parvis compositi, cellulas conidiogenas apicales vel laterales ferentes, ramis secundaris raris. Cellulae conidiogenae cylindricae, polyblasticae, integratae, sympodiales, cicatricatae. Conidia 12–25 (18) x 3–5 (4) µm, pallido brunnea vel brunnea, laevia, acropleurogena, solitaria, elliptica, 1–2-septata, aliquando cum constrictionibus septalibus, cum basibus truncatis.
Colonies hypophyllous, grayish, velutinous, effuse, growing among leaf ribs. Mycelium superficial or partially immersed. Hyphae septate, branched, hyaline to pale brown, smooth. Conidiophores 250–520 (320) x 7–11 (9) µm, macronematous, mononematous, straight, seldom curved, brown to dark brown, septate, smooth, with primary branches located on the apical one-third of the stipe (Fig. 33). Primary branches 23–69 (43) x 10 µm, 2–6-small celled, with lateral and apical conidiogenous cells, secondary branches rare (Figs. 34–36). Conidiogenous cells cylindrical, polyblastic, integrated, sympodial, cicatrized. Conidia 12–25 (18) x 3–5 (4) µm, pale brown to brown, smooth, acropleurogenous, single, elliptical, 1–2-septate, sometimes with constriction at the septa, basally truncate (Fig. 35).
Specimens examined. BRAZIL. DF: Brasília, Parque Nacional de Brasília, on living leaves of Qualea grandiflora, 21 Aug 1995, MF Almeida n. 13 (HOLOTYPUS:UB col. micol. 9851). BRAZIL. DF: Brasília, Parque Nacional de Brasília, on Qualea grandiflora, 21 Aug 1995, Z M Chaves n. 143 (UB col. micol. 9817).
This fungus fits the concept of Periconiella (Ellis 1967) and shows a typical feature—the apical primary branches originate from the most apical down to the sixth most apical cells of the conidiophore axis; such branching may occupy up to one third of the stipe. Species having 2–3 celled solitary conidia are limited to P. phormii Ellis and P. telopeae (Hansf.) Ellis (Ellis 1967, 1971, 1976). Periconiella qualeae-grandiflorae differs from both because of a much broader extension of the branched portion of its conidiophore and smooth instead of verrucose conidial walls. Thus this fungus from Q. grandiflora belongs in a new taxon, P. qualeae-grandiflorae.
Periconiella campo-grandensis Dornelo-Silva & Dianese, sp. nov. Figs. 37–42
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Coloniae saepe in trichomate insidentes, hypophyllae, cinerascentes, velutinae, intercostales. Mycelium superficiale ex parte immersum. Hyphae septatae, ramosae, hyalinae vel pallido brunneae, laeviae. Conidiophora 150–370 (235) x 5–6 (5) µm, macronemata, mononemata, erecta, aliquando curvata, raro ramosa, brunnea vel atro brunnea, septata, laevia, ramis primariis et secundaris capitulo apicali formantibus. Rami primarii 15–36 x 5–6 (5) µm. Rami secundarii 16–20 x 4–5 µm. Cellulae conidiogenae cylindricae, laeviae vel verruculosa, polyblasticae, integratae, sympodiales, cicatricatae. Conidia 7–15 (10) x 4–6 (5) µm, acropleurogena, solitaria, elliptica, hyalina vel pallido brunnea, aseptata, verrucosa, cum basibus truncatis et cicatricatis.
Colonies mostly on trichomes (Figs. 37, 38), hypophyllous, grayish, velutinous, located among leaf ribs. Mycelium superficial, partially immersed. Hyphae septate, branched, hyaline to pale brown, smooth. Conidiophores 150–370 (235) x 5–6 (5) µm, macronematous, mononematous, straight, seldom curved, brown to dark brown, septate, smooth, sometimes branched near the middle of the stipe, with primary and secondary conidiogenous branches forming an apical head (Figs. 37–40). Primary branches 15–36 x 5–6 (5) µm (Figs. 39, 40). Secondary branches 16–20 x 4–5 µm (Figs. 40, 41). Conidiogenous cells cylindrical, smooth or verruculose, polyblastic, integrated, sympodial, cicatrized (Figs. 40, 41). Conidia 7–15 (10) x 4–6 (5) µm, acropleurogenous, single, elliptical, hyaline to pale brown, aseptate, verrucose, with base truncate and cicatrized (Fig. 42).
Specimens examined. BRAZIL. MS: Campo Grande, Fazenda Lagoinha-Embrapa, on living leaves of Qualea grandiflora, 19 Aug 1996, CA Inacio n. 582 (HOLOTYPUS: UB col. micol. 12459). BRAZIL. DF: Planaltina, Estação Ecológica das Águas Emendadas, on Qualea grandiflora, 13 Jun 1995, ZM Chaves n. 100 (UB col. micol. 8787).
Among known Periconiella species, only P. angusiana Ellis shows conidiophores with primary and secondary branches and aseptate verrucose conidia (Ellis 1967). However, P. angusiana has conidiophores with verrucose primary, secondary and tertiary branches and not smooth primary and secondary branches, as occurs in P. campo-grandensis.
The three new Periconiella species described on Q. grandiflora can be identified using this key:
KEY OF SPECIES OF PERICONIELLA ON Q. GRANDIFLORA
1. Conidia aseptate, verrucose, 7–15 µm long,, conidiophores 150–370 µm long, primary branches 15–36 µm long . . . . . P. campo-grandensis sp. nov.
1. Conidia septate, smooth . . . . . 2
2. Conidia 1–2-septate, elliptical, clavate or obclavate; 12–25 µm long, conidiophores up to 520 µm long with primary branches 23–69 µm long . . . . . P. qualeae-grandiflorae sp. nov.
2. Conidia 3–5-septate, cylindrical; 37–95 µm long, condiophores up to 770 µm long with primary branches 60–137 µm long . . . . . P. longispora sp. nov.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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Accepted for publication April 7, 2003.
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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———. 1994b Alternaria themes and variations (106–111). Mycotaxon 50:409-427
———. 1997 Alternaria themes and variations (151–223). Mycotaxon 65:1-91
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