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Mycologia, 96(3), 2004, pp. 675-681.
© 2004 by The Mycological Society of America

Poroleprieuria, a new xylariaceous genus from Mexico


María C. González 1

     Laboratorio de Micromicetes, Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Ciudad de México, D.F., 04510, México

Richard T. Hanlin

     Department of Plant Pathology, University of Georgia, Athens, Georgia 30602-7274

Miguel Ulloa
Elvira Aguirre

     Laboratorio de Micromicetes, Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Ciudad de México, D.F., 04510, México

   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
 LITERATURE CITED
 

Poroleprieuria gen. nov. is described and illustrated to accommodate P. rogersii in the Xylariaceae, Xylariales. This ascomycete, known only from the type collection, is characterized by reniform, light brown, smooth ascospores with a germ pore; cylindrical, persistent asci lacking an apical apparatus, septate persistent paraphyses, and erumpent, erect, dark brown, fragile, subcylindrical stromata. The characteristics of this xylariaceous fungus were compared with those of some other ascomycetes having superficially similar cylindrical stromata or ascospores with germ pores.

Key words: Ascomycota, Leprieuria, lignicolous, systematics, Xylariaceae, Xylariales


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
 LITERATURE CITED
 
The Xylariales is the best-studied group of ascomycetes from Mexico (Lodge 2001Go). The first work was performed by Fries (1851)Go, and later several additional studies were undertaken (Saccardo 1891Go, Sharp 1948Go, Welden and Lemke 1961Go, Guzmán and Madrigal 1967Go, Pérez-Silva 1975Go). However, the first continuous systematic research on xylariaceous fungi was initiated in this country by San Martín and Rogers (1989)Go, which has resulted in several publications on the Xylariaceae (San Martín and Rogers 1993aGo, bGo; Ju et al 1998Go; San Martín and Lavín 1999Go; San Martín et al 1997Go, 2001Go; Rogers et al 2002aGo).

During the relocation of the old Instituto de Biología building to a new location, an unusual xylariaceous specimen was found in the fungal collection of the Herbario Nacional de México. This interesting fungus has combined characteristics of reniform, light brown, smooth ascospores with a germ pore; cylindrical persistent asci; ascus tip inamyloid; persistent paraphyses; and erumpent, erect, cylindrical stromata. These characteristics place the fungus in order Xylariales. Comparison with published descriptions indicates that it is an undescribed genus. The relationships of this Mexican ascomycete with Leprieuria and other genera of family Xylariaceae is discussed.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
TAXONOMY
 DISCUSSION
 LITERATURE CITED
 
The fungus was collected near Ayotoxco de Guerrero City, in Ayotoxco de Guerrero municipality, state of Puebla, in the central part of Mexico (FIG. 1Go). It was sent to M. Ulloa by the collectors in Aug 1989, and the specimen was placed in the fungi collection of Herbario Nacional de México (MEXU) of the Universidad Nacional Autónoma de México and also was deposited in the J.H. Miller Mycological Herbarium (GAM) of the University of Georgia. During a reorganization of the mycology collection, the specimen recently was rediscovered. The state of Puebla has an area of 33 995 km2 and includes 217 municipalities. The Ayotoxco de Guerrero municipality has an area of 130 km2 and is located in northeastern Puebla (between 19°59'54''N, 97°21'18''W and 20°08'48''N, 97°27'42''W), its altitude varies between 100 and 520 m, its climate is hot and humid with rain throughout the year, the annual median temperature is 22 C, precipitation during the driest month is 60 mm; the land is covered with pastures, small areas of coffee plantations and perennial forest associated with secondary shrub vegetation (INEGI 2000Go).



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  		FIG. 1. Map of Mexico showing Ayotoxco de Guerrero municipality of the state of Puebla, in which the fungus was discovered.

 
Squash mounts of the fungus for observations of structures were made in water. A drop of Melzer’s reagent was added to determine the amyloid iodine reaction, or lack of a reaction, of the ascal apex. In addition, the structures were examined in lactophenol and stained with cotton blue. Measurements of 25 structures (stromata, ascomata, paraphyses, asci and ascospores) were made on rehydrated material mounted in water. Four types of microscopy were used in this study: light microscopy (LM), differential interference contrast microscopy (DICM), phase contrast microscopy (PHCM) and scanning electron microscopy (SEM). Ascospores examined by SEM were air dried and coated with gold palladium. Attempts were made to culture the fungus, but in all cases were unsuccessful.


   TAXONOMY
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
DISCUSSION
 LITERATURE CITED
 
Poroleprieuria M. C. González, Hanlin, Ulloa et E. Aguirre, gen. nov.

Stromata per corticem erumpentia, erecta, subcylindrica, laete brunnea, fragilia. Ascomata ovoideoelongata, in stromatis parte supera contigue immersa. Asci clavati, octospori, leptodermi, unitunicati, persistentes. Paraphyses filamentosae, septatae, persistentes. Ascosporae reniformes, unicellulares, pallido-brunneae, exappendiculatae, laeves, poro germinali apicali.

Stromata erumpent through bark, erect, cylindrical, dark brown, fragile. Ascomata elongate-ovoid, packed in the upper part of the stroma. Asci clavate, 8-spored, thin-walled, unitunicate, persistent. Paraphyses filamentous, septate, persistent. Ascospores reniform, 1-celled, light-brown, not appendaged, smooth, with an apical germ pore.

Etymology. – From the L. porus = passage, in reference to the germ pore of the ascospores + leprieuria, named in honor of C. Leprieur, a collector of Xylariaceae in French Guiana.

Type species. – Poroleprieuria rogersii.

Poroleprieuria rogersii M. C. González, Hanlin, Ulloa et E. Aguirre, sp. nov. FIGS. 2Go–24Go



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  		FIGS. 2–12. Poroleprieuria rogersii. 2. Lateral view of stroma showing the superficial annulations. 3. Stromata erumpent through bark. 4. Stromata apex showing the ostioles. 5. Ostioles. 6. Stroma apex. 7. Ostioles lower than the stromatal surface. 8. Top view of stroma with ostioles filled with ascospores. 9. Transverse section of the stroma showing the endostroma and ascomata centrum. 10. Transverse section revealing the ascoma wall. 11. Ascoma wall. 12. Ascoma wall with ascogenous system that lines the ascoma wall. Scale bars: 2–4 = 1000 µm, 5 = 25 µm, 6, 7 = 200 µm, 8, 9 = 250 µm, 10 = 100 µm, 11 = 20 µm, 12 = 8 µm. Micrographs from HOLOTYPE, LM: 2–4, SEM: 5–12.

 


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  		FIGS. 13–24. Poroleprieuria rogersii. 13. Longitudinal section of stroma showing the ascomata. 14. Longitudinal section of stroma showing the endostroma and the ascogenous system that lines the ascoma wall. 15. Cluster of asci and ascospores. 16. Branched paraphyses. 17. Young ascus with eight ascospores. 18. Ascospores. 19. Group of eight young ascospores. 20. Ascospores with germ pore. 21. Ascospore terminal germ pore. 22, 23. Ascospores with terminal gem pore. 24. Detailed ascospore germ pore. Scale bars: 13 = 500 µm, 14 = 20 µm, 15 = 40 µm, 16 = 20 µm, 17 = 12 µm, 18 = 8 µm, 19 = 3.5 µm, 20 = 5 µm, 21, 22 = 1 µm, 23 = 1 µm, 24 = 500 nm. Micrographs from HOLOTYPE, SEM: 13, 14, 19–24, DICM: 15, PHCM: 16, 17, LM: 18.

 
Stromata per corticem erumpentia, gregaria, erecta, sub-cylindrica, attenuata (1750–)5750–6750(–7750) x (550–)700–850(–1050) µm diametro ad apicem, basi (450–)650–700 (–1050) µm diametro, longitudinaliter sulcata, dimidio inferiori annulato, ad basem per annulum brevem volvifor-men circumcincta, annulo 50–750 alto x 100–250 µm cras-so constato ex contexto ectostromatis hospitisque, apicem leviter clavato, convexa, 2–10 ostiolis, singulatim erumpen-tibus, quam pagina stromatali umbilicata inferioribus, centrale dispositis, solida ad apicem, basi excavata. Ectostroma tenue, fragile, atrobrunneum. Endostroma ad apicem soli-dum, atrobrunneum, et parte media molle, fibrosum, albi-dum vel eburneum ad basem. Ascomata ovoideo-elongata, 806–1484 x 161–364 µm, immersa, pariete tenui 4–8 µm crassa, 2–10 ascomata per stroma, in entostroma atro-brun-neo contigua, apicem insidentia, singulatim erumpentia. Asci cylindrici, 71.5–82 x 3.5–6 µm, apice rotundati, brevistipitati, octospori, persistentes, leptodermi, unitunicati, monostichi, sine apparato apicali, nonamyloidei, constati ex systema ascogeno parietem ascomatalem tegenti. Paraphyses filiformes, 140–164 x 3–4 µm, 3 µm latae, septatae, septo leviter constricto, persistentes, leptodermae, ramosae prope basin. Ascosporae reniformes, pallido-brunneae, (6–)7–8(–9) x (3–)4–4.5(–5) µm unicellulares, uniseriatae, exappen-diculatae, sub LM, DICM et SEM laeves, poro germinali prominenti terminali. Anamorpha: ignota.

Stromata erumpent through bark, gregarious, erect, subcylindrical, slightly tapered (1750–)5750–6750(–7750) x (550–)700–850(–1050) µm at the apex and (450–)650–700(–1050) µm at the base, longitudinally sulcate over the entire length, annulate on the lower half, basally surrounded by a short volva-like ring 50–750 high x 100–250 µm wide, composed of ectostromatal and host tissue; apex slightly clavate, convex, with 2–10 ostioles individually erumpent, lower than the stromatal surface (umbilicate), centrally disposed, solid in the apical part and hollow in the basal part. Ectostroma thin, brittle, fragile, dark brown. Endostroma solid and dark brown in the apical part, and soft, fibrous, white to tan in the middle and basal part. Ascomata elongate-ovoid, 806–1484 x 161–364 µm, with a thin wall 4–8 µm thick, 2–10 ascomata per stroma, packed in a dark brown endostroma, apically seated, individually erumpent. Asci cylindrical, 71.5–82 x 3.5–6.0 µm, rounded at the top, short stipitate, 8-spored, persistent, thin-walled, unitunicate, monostichous, apical apparatus lacking, inamyloid, formed from an ascogenous system that lines the ascomatal wall. Paraphyses filamentous, 140–164 x 3–4 µm, septate, slightly constricted at the septa, persistent, thin walled, branched in the basal part. Ascospores reniform, light brown, (6–)7–8(–9) x (3–)4–4.5(–5) µm, 1-celled, uniseriate, no appendage, spore wall smooth as seen by LM, DICM, and SEM, with one prominent terminal germ pore.

Anamorph: unknown.

Specimen examined. – MEXICO. STATE OF PUEBLA: Ayotoxco de Guerrero municipality, 4 km East Rancho La Carolina. On decayed bark of Heliocarpus sp. (Tiliaceae), 25 Aug 1989, W. López-Forment and A. Ocampo (HOLOTY PE MEXU 23019, ISOTY PE GAM 16055).

Etymology. – The epithet rogersii refers to the last name of Jack David Rogers, an outstanding mycologist, who has contributed greatly to our knowledge of the xylariaceous fungi.


   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
LITERATURE CITED
 
The taxonomic affinities of Poroleprieuria are unclear because it has a set of unusual morphological attributes (Rogers 1994Go). It is characterized by its erect, cylindrical, dark-colored stroma, with ascomata embedded in the upper part, near the apex; cylindrical, persistent asci, without an apical apparatus; reniform, light brown, 1-celled, smooth walled ascospores, with one terminal germ pore. The lignicolous habit and the morphological characteristics of Poroleprieuria are unlike those of any described fungus, but they suggest some linkage to the Xylariales. The genus that exhibits the greatest morphological similarity to Poroleprieuria is Leprieuria Læssøe, J.D. Rogers & Whalley of the Xylariaceae (Xylariales) (Læssøe 1994Go). The genera Camarops P. Karst (Boliniaceae, Boliniales), Sordaria Ces. & De Not. (Sordariaceae, Sordariales) are discussed briefly because the species of these genera have 1-celled ascospores with germ pores and therefore possibly could be related to Poroleprieuria. In addition, the genus Camillea Fr. is discussed on the basis of its erect stromata, ascospore color and lignicolous habit. Because these differ in some significant ways and probably are unrelated, species of these genera exhibiting ascospores with germ pores were not considered further: Apiorhyn-chostoma Petr.; Ommatomyces Kohlm. Volkm.-Kohlm & O.E. Erikss. (Clypeosphaeriaceae, Xylariales); Apiocamarops Samuels & J.D. Rogers; and Pseudovalsaria Spooner (Boliniaceae, Boliniales).

The monotypic genus Leprieuria, erected by Læssøe et al (1989)Go, is considered to include xylariaceous fungi with erect, cylindrical, black stromata with reniform, dark brown ascospores with prominent, ventrally situated, straight, germ slits. They excluded this taxon from Camillea Fr. because of the presence of a Geniculosporium Chesters & Greenh. anamorph and ascospores lacking ornamentations that have ventrally situated straight germ slits (Læssøe et al 1989Go). Leprieuria bacillum (Mont.) Læssøe, J.D. Rogers & Whalley is fundamentally similar to the type species of Poroleprieuria. Although both species are similar in the morphology of the stromata and ascospores, L. bacillum differs from P. rogersii in that it has a black stroma of a different size (3000–6000 x 500–750 µm) with a smaller number of ostioles (2–6); deliquescent asci and paraphyses; and dark brown to black ascospores with prominent, ventrally situated straight germ slits of a smaller size (5.5–7 x 3–4 µm) (Montagne 1840Go, Dennis 1957Go, Samuels and Müller 1980Go); and a Geniculosporium anamorph. Poroleprieuria is not known to have an anamorph. Based on the above, it is not likely that these taxa are congeneric.

Camarops P. Karst. long was considered to be related to the Xylariaceae, but molecular data based on 18S rDNA sequence data indicates a closer link with the Sordariales, and presently it is placed in the Boliniaceae (Andersson et al 1995Go). Camarops is similar to Poroleprieuria because it has unicellular brown ascospores, with apical germ pores, persistent asci with non-amyloid apex, stromata and lack of anamorphs (Nannfeldt 1972Go). However, the differences between these genera suggest that they are not related taxa. The ascospores of Camarops are ellipsoidal, usually flattened and bearing a germ pore at one or both ends, minute (5 x 2–3 µm); asci are small; stromata are variable, but none are cylindrical.

Sordaria Ces. & De Not. is characterized by dark, nonstromatic ascomata; cylindrical asci with a distinct nonamyloid apical ring, and dark brown, ovoid, 1-celled, smooth ascospores with a basal germ pore that are covered with a gelatinous sheath (Lundqvist 1972Go). Poroleprieuria differs from Sordaria in having stromatic ascomata, asci lacking an apical apparatus, and reniform, light brown ascospores without gelatinous sheaths. The Sordariaceae in the morphology of the ascospore and ascus apex resembles the Xylariaceae, but the predominance of phialidic conidia, ascospore germ pores and nonstromatic ascocarps indicates lack of a close relationship to the Xylariaceae (Rogers 1979Go). Poroleprieuria interascal tissue is composed of persistent, wide, thin-walled, basally branched paraphyses. Sordaria humana (Fuckel) Winter has persistent wide paraphyses with thin cell walls and irregular branching (Read and Beckett 1984Go). The morphological similarities between Poroleprieuria and Sordaria indicate a possible relationship, but it remains obscure until new data are obtained.

Camillea Fr. is a genus with ascospores atypical of the Xylariaceae. Light-colored ascospores of Camillea are ornamented variously with pits, spines or much more complicated structures (Rogers 1977Go), and without germ slits, except for one species (Rogers et al 2002bGo). In addition, Camillea has bipartite stromata that are applanate to erect, tall or short, with apices mucronate, bilabiate, depressed-discoid or pla-no-convex, and the anamorph is Xylocladium P. Syd. (Læssøe et al 1989Go). Poroleprieuria has a cylindrical stroma, reminiscent of Camillea leprieurii Mont., but other characters differ significantly. The morphological similarity between the erect form of Camillea leprieurii and Poroleprieuria could be the result of convergent evolution.

The stromata of xylariaceous fungi function in part to protect perithecia from desiccation and appear to be adaptations to minimize dehydration in dry environments (Rogers 1979Go). The new taxon was discovered in a tropical region of the state of Puebla. Climate is an important factor affecting the distribution of Xylariaceae; however, the climatic zone might not be the only factor to consider in the distribution (Rogers 2000Go, Whalley 1985Go). Xylariaceous genera with stalked stromata raise their ascomata above the substrate thus effecting more efficient ascospore dissemination (Whalley 1985Go). The dispersal of P. rogersii is unknown but probably is similar to other taxa with erect stromata. Ascospores are particularly subject to environmental selection and are consequently of value at lower taxonomic ranks (Hawksworth and Mouchacca 1994Go). Ascospore morphology apparently is not much affected by biotic and abiotic factors. Size, shape, color and surface ornamentation all are useful characters in the delimitation of taxa. However, the usefulness of ascospore characters depends upon the degree of correlation with other characters of the stromata, ascomata, asci, anamorph and habitat (Rogers 1979Go). Poroleprieuria ascospores are devoid of a germ slit, but this taxon shows other cardinal features of the Xylariaceae, such as stromata, ascomata, asci, ascospore shape and habitat. Anamorph characteristics of P. rogersii unfortunately were not revealed, despite several attempts to culture the fungus. There are a number of xylariaceous fungi with ascospores that apparently are devoid of germ slits or other germination sites, and ascospores lacking the slit might have germination sites in the wall that are not obvious. Ascospores devoid of the slit, in general, are lighter in color than those having it (Rogers 1979Go). Poroleprieuria features light-colored poroid ascospores with the occurrence of ascospores without evident germ pores, possibly because of a lack of contrast between the pale wall and the orifice.


   ACKNOWLEDGMENTS
 
The authors are grateful to Carmen Rodríguez and Berenit Mendoza for their technical assistance with SEM. We are grateful to Calixto León for determining the host-plant species. We also thank Fernando Chiang for correcting the Latin diagnosis. This study was supported by the Instituto de Biología, Universidad Nacional Autónoma de México, and the Department of Plant Pathology, University of Georgia.


   FOOTNOTES
 
Accepted for publication September 25, 2003.

1 Corresponding author. E-mail: mcgv{at}ibiologia.unam.mx


   LITERATURE CITED
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 TAXONOMY
 DISCUSSION
 LITERATURE CITED
 
Andersson K, Eriksson OE, Landvik S. 1995. Boliniaceae transferred to Sordariales (Ascomycota). Syst Ascomycetum 14:1–16.

Dennis RWG. 1957. Further notes on tropical American Xylariaceae. Kew Bull 2:297–333.

Fries EM. 1851. [New mycological contributions of foreign lands from collections made by Danish botanists.] Nova Acta Reg Soc Sci Uppsal Ser 3, 1:1–136. [In Latin]

Guzmán G, Madrigal X. 1967. [Notes on some higher fungi from Escárcega, Campeche, Mexico.] Rev Soc Mex Hist Nat 28:23–38. [In Spanish]

Hawksworth DL, Mouchacca J. 1994. Ascomycete systematics in the nineties. In: Hawksworth DL, ed. Ascomycete Systematics. Problems and perspectives in the nineties. New York: Plenum Press. p 3–11.

[INEGI] Instituto Nacional de Estadística, Geografía e Informática. 2000. [Geographical abstract of the state of Puebla.] Aguascalientes, México: Instituto Nacional de Estadística, Geografía e Informática. 768 p. [In Spanish]

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Læssøe T. 1994. Index ascomycetum I. Xylariaceae. Syst Ascomycetum 13:43–112.

———, Rogers JD, Whalley JS. 1989. Camillea, Jongiella and light-spored species of Hypoxylon. Mycol Res 93:121–155.

Lodge JD. 2001. World and regional diversity of fungi. In: Hernández HM, García-Aldrete AN, Álvarez F, Ulloa M, eds. Contemporary approaches for the study of biodiversity. México: Instituto de Biología, UNAM. p 291–304. [In Spanish]

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———. 1979. The Xylariaceae: systematic, biological and evolutionary aspects. Mycologia 71:1–42.

———. 1994. Problem genera and family interfaces in the Eupyrenomycetes. In: Hawksworth DL, ed. Ascomycete Systematics. Problems and perspectives in the nineties. New York: Plenum Press. p 321–331.

———. 2000. Thoughts and musings on tropical Xylariaceae. Mycol Res 104:1412–1420.

———, San Martín F, Ju YM. 2002a. A reassessment of the Xylaria on Liquidambar fruits and two new taxa on Magnolia fruits. Sydowia 54:91–97.

———, ———, ———. 2002b. Three new taxa of Camillea from Costa Rica. Sydowia 54:84–90.

Saccardo PA. 1891. Sylloge Fungorum. Pavia. Vol. 9.

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