ARTÍCULOS ORIGINALES

 

Biodiversity of Xylariaceae (Ascomycota) and their hosts in protected areas from Tucuman (Argentina)

Biodiversidad de la familia Xylariaceae (Ascomycota) y sus hospedantes en áreas protegidas de Tucumán (Argentina)

 

Esteban B. Sir ^{1 2}*, Adriana I. Hladki ¹, Maria F. Parrado ³ & Andrea I. Romero ⁴

¹ Laboratorio Criptogámico, Fundación Miguel Lillo. Miguel Lillo 251, San Miguel de Tucumán (CP 4000), Tucumán, Argentina.
² CONICET.
³ Laboratorio de Taxonomía Vegetal, Fundación Miguel Lillo.
⁴ PRHIDEB-CONICET, Dep. de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales (UBA), Ciudad Universitaria, Pabellón II, 4to. Pio, CP1428EHA Buenos Aires, Argentina.
* Corresponding author: Sir E. B. sirestebanbenjamin@gmail.com

 

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Abstract

Fourty nine species belonging to the genera Astrocystis, Biscogniauxia, Daldinia, Entonaema, Hypoxylon, Kretzschmaria, Phylacia, Poronia, Rosellinia, Stilbohypoxylon and Xylaria were recorded in protected areas from Tucuman. The species are grouped in probably endemic, cosmopolitan, neotropical and pantropical. The biodiversity of Xylariaceae in this province is compared with those from other countries. Twenty-three host taxa are registered, most of them were dicots. Cinnamomum porphyrium and Podocarpus parlatorei were the two native hosts more colonized.

Key words: Argentina; Mycogeography; Xylariales; "Yungas".

Resumen

Se registran cuarenta y nueve especies pertenecientes a los géneros Astrocystis, Biscogniauxia, Daldinia, Entonaema, Hypoxylon, Kretzschmaria, Phylacia, Poronia, Rosellinia, Stilbohypoxylon y Xylaria. Las especies se agrupan, según su distribución, en endémica probablemente, cosmopolita, neotropical y pantropical. Se compara la biodiversidad de Xylariales de la provincia de Tucumán con la de otros países. Se identifican veintitrés hospedantes, la mayoría de ellos dicotiledóneas, siendo Cinnamomum porphyrium y Podocarpus parlatorei las especies nativas más colonizadas.

Palabras clave: Argentina; Micogeografía; Xylariales, "Yungas".

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Introduction

Xylariaceae is the largest family of the order Xylariales and one of the largest families in Ascomycota. Although its distribution is cosmopolitan, its greatest diversity is found in the tropical and subtropical regions.

The knowledge on the family in the South Cone of South America is largely limited to the contributions made by Spegazzini (1880a, b, c, 1881, 1884, 1887, 1888a, b, 1889, 1891, 1898, 1899, 1902, 1908a, b, 1909, 1910, 1912, 1919, 1921, 1922). Part of his work is devoted to Xylariales of Argentina, Bolivia, Brazil, Chile and Paraguay. Later Theissen (1908, 1909a, b), Rick (1935), Chardon et al. (1940) described many species from Brazil and Dennis (1956, 1957, 1958, 1970) helped to clarify the nomenclatural and taxonomic status of the South American tropical Xylariales. Rogers et al. (1988) contributed valuable information to Venezuela, Laessoe (2002) conducted a survey of the family in Ecuador and new data for this country can be seen in http://www.mycokey.com/Ecuador.html. Hamme (1993), Hamme & Guerrero (1997, 2002), Pereira et. al (2008) and Trierveiler-Pereira et al. (2009) recently investigated presence of Xylaria Hill ex Schrank and Kretzschmaria Fr. in Brazil.

A total of 84 species were reported by Spegazzini (citations mentioned above) of the following genera from Argentina: Anthostomella Sacc. (11 spp.), Daldinia Ces. & De Not. (5 spp.), Hypocopra (Fr.) J. Kickx. f. (6 spp.), Hypoxylon Bull. (19 spp.), Kretzschmaria (2 spp.), Poronia Willd. (3 spp.), Rosellinia De Not. (13 spp.) and Xylaria (25 spp.). From Tucumán province Hypoxylon notatum (Hypoxylon nectrioide Speg.), Roselliniaparaguayensis Starbäck, R. smilacina Speg., Xylaria arbuscula Sacc., X. gomphus Fr., X. ianthinovelutina (Mont.) Fr., X. macropoda Speg., X. smilacicola Speg., X. sordida Speg., X. torulosa (Jungh.) Speg. and X. venosula Speg. were recorded. Dennis (1956, 1957, 1958) reported 14 species of the following genera from Tucuman: Xylaria (9 spp.), Poronia (2 spp.), Phylacia Lev. (1 sp.), withX. ianthinovelutina, X. grammica (Mont.) Mont., X. smilacicola, Poronia punctata (L.) Br. and X telfairii (Berk.) Sacc.

In order to know more about the biodiversity of the xylariaceous in Tucuman and their hosts, numerous investigations were carried out (Hladki 1997, 2001, 2004; Hladki & Romero 2001, 2003, 2005, 2006, 2007, 2009a, b, 2010; Sir et al. 2012). As a consequence of all the studies of the genera and species that have been carried out and are provided here as one of the aims of this article, we have been able to compare and discuss the composition of the local mycobiota with others from tropical, subtropical and temperate regions and the relation between each xylariaceous species and their identified hosts.

Materials and methods

Thirty seasonal mycological expeditions in protected areas of the Tucumán province (1-Provincial Park "The Cochuna", 1306 m. s.l.; 2-Reserva Provincial La Florida, 429 m.s.l.; 3-"Biological Park" Sierra de San Javier", 1153 m.s.l.) (Fig. 1) were made (Details in Hladki 2001, 2004; Hladki & Romero 2001, 2003, 2005, 2007, 2009b, 2010). Types and other collections from South America kept at BPI, C, FH, GZU, ICN, IMI, K, LPS, NY, PC, S, W, WSP (Thiers 2012) have been examined. In total 961 collections were gathered from the Tucumán province (LIL). Microscopic preparations and observations have been performed according to Ju & Rogers (1996). The results are organized in the following sections:

Fig. 1. Map "Las Yungas", Sampling Sites: 1, Provincial Park "The Cochuna". 2, Reserva Provincial La Florida. 3, "Biological Park" Sierra de San Javier.

Composition of xylariaceous mycobiota in Tucumán province

The total number of the identified species is shown and listed. In addition, the species were classified into groups according to the distributional patterns (Cosmopolitan, probably Endemic from Tucuman Yungas (YUT), Neotropical, Pantropical) and their percentages are shown graphically.

Local mycobiota compared with others from other regions

The local xylariaceous taxa are compared with other areas of different countries from neotropical, paleotropical and holartic kingdoms (Takhtajan 1986) through the species richness (n) per genus. The compared areas are arranged according to the size expressed in km², from the smallest (Tucuman) to largest (Europe); taking areas into account where exhaustive studied on the family were carried out independently if the area was a state, a country or a continent. Data for each country (from each studied area) were compiled from the reviewed literature. A total of 50 publications including papers, books, monographs and web pages, in which xylariaceous fungi have been cited were reviewed (Bitzer et al. 2007; Camino Vilaró et al. 2006; Carmona et al. 2009; Dennis 1961, 1963, 1970; Francis 1975; Granmo et al. 1999; Hellwing et al. 2005; Ju, & Rogers 1999; Lee & Crous 2003; Lu & Hyde 2000; Medel et al. 1999, 2008; Meijer 2006; Mühlbauer et al. 2002; Okane et al. 2008; Pereira et al. 2010; Pérez-Silva 1972, 1975; Poroca 1986; Quang et al. 2005; Rogers et al. 1987, 1988; Rogers & Ju 2012; San Martín & Lavin 1997; San Martin & Rogers 1989, 1993, 1995; San Martin et al. 1997; Stadler & Baumgartner 2001; Stadler et al. 2001a, b, c; 2002; 2004a, b, c; 2007; Taligoola & Whalley 1976, 1977; Triebel et al. 2005;Van der Gucht 1995, 1996; Vasilyeva & Stadler 2008; Whalley et al. 1988; Wollweber & Stadler 2001; www.mykokey.com/ecuador; http//:www.cybertruffle.org.uk; http://mycology.sinica.edu.tw/Xylariaceae, http://pyrenomycetes.free.fr). All these data are displayed through a table and are expressed graphically.

Hosts

The biogeographic province, "Las Yungas", has a subtropical climate and belong to the "Amazonic Dominion" (Phytogeographical region according to Cabrera & Willink (1980), being a Neotropical region. "Las Yungas" is distributed parallel to the Andes from Venezuela and Colombia to Catamarca state in Argentina (Cabrera & Willink, 1980; Hueck 1978). It is integrated by elements from the "Amazonic Dominion" -Inga Mill., Nectandra Rottboell, Ocotea Aublet, Citronella D. Don, Cedrela P. Br., Phoebe Ness, Miconia R. et P. -and being placed in the Andean corridor also has elements from Holartic -Viburnum L., Juglans L., Rhamnus L., Myrica L., Alnus Mill., Berberís L., Rubus L., Sambucus L., Prunus L., Ilex L. -and "Gondwanic" -Podocarpus L'Hér., Gunnera L., Fuchsia L., etc. -origin and also others elements in common with the south American mountain forest -Oreopanax Decaisne et Planchon, Tabebuia DC., Crinodendron Molina, Chusquea Kunth-(Brown 1995).

In Argentina, "Las Yungas" covers a narrow strip, 2.700.000 hectares (23-29° S, 64-68° O) North-South, along Salta, Jujuy, Tucumán and Catamarca provinces, between 300 and 3000 m at above sea level (s.l.m.). Although "Las Yungas" represents a marginal ecosystem for Argentina (less of 0.1% of the territory) it has almost 50% of the Argentine biodiversity (Brown 1995; Brown et al. 1985; Brown & Ramadori, 1989; Brown et al. 2002).

The main plants of the Tucuman Yungas in the lowest altitudes (between 400 to 700 m.s.l.) are Calycophyllum multiflorum (Rubiaceae), Phyllostylon rhamnoides (Ulmaceae), Handroanthus impetiginosus (Bignoniaceae), among others (Brown 1995). In the range of 700 to 1500 m.s.l., it is dominated by Ficus maroma (Moraceae), Alnus acuminata (Betulaceae), Cinnamomum porphyrium, Nectandra pichurim and Ocotea puberula (Lauraceae), Inga edulis, I. semialata, I. saltensis y Tipuana tipu (Fabaceae) and Blepharocalix salicifolius (Myrtaceae). Finally up to 3000 m.s.l. it is found the Podocarpus parlatorei (Podocarpaceae) forest, Roupala meisneri (Proteaceae) and Fuchsia boliviana (Onagraceae), Alnus acuminata (Betulaceae) among others (Brown et al. 2002)

The relationship between each species and the type of hosts (gymnosperms; angiosperms: monocots, dicots) will be shown through a table and the percentages of colonized hosts by the xylariaceous taxa are graphically presented.

Results

Composition of xylariaceous mycobiota

We recognized 49 taxa in protected areas of the province of Tucumán: Astrocystis smilacicola (Schwein.) Laessoe & Spooner, Biscogniauxia capnodes (Berk.) Y. M. Ju & J. D. Rogers, Daldinia eschscholzii (Ehrenb.) Rehm, Entonaema liquescens Möller, Hypoxylon anthochroum Berk. & Broome, H. notatum Berk. & M. A. Curtis, H. rubiginosum var. microsporum Whalley, H. subrutilum Starbäck, Kretzschmaria argentinensis Hladki & A. I. Romero, K. clavus (Fr.) Sacc., K. deusta (Hoffm.) P. M. D. Martin, K. pavimentosa (Ces.) P. M. D. Martin, K. sandvicensis (Reichardt) J. D. Rogers & Y. M. Ju, K. sigmoidirima Hladki & A. I. Romero, Phylacia globosa Lév., Poronia oedipus (Mont.) Mont., P. punctata, Rosellinia necatrix Berl. ex Prill., R. paraguayensis, R. subiculata (Schwein.) Sacc., Stilbohypoxylon macrosporum Hladki & A. I. Romero, S. minus Hladki & A. I. Romero, Xylaria adscendens (Fr.) Fr., X. allantoidea (Berk.) Fr. , X. apiculata Cooke, X. coccophora Mont., X. cubensis (Mont.) Fr., X. culicicephala A. I. Romero & Hladki, X. curta Fr., X. enteroleuca (Speg.) P. M. D. Martin, X. filiformioidea Hladki & A. I. Romero, X. fissilis Ces., X. globosa (Spreng. ex Fr.) Mont., X. gracillima (Fr.) Fr., X. grammica, X. ianthinovelutina, X. aff. ianthinovelutina, X. kretzschmarioidea J. D. Rogers & Rossman, X. luxurians (Rehm) Lloyd, X. melanura Lév., X. mellissii (Berk.) Cooke, X. microceras var. yungae Hladki & A. I. Romero, X. multiplex (Kunze) Fr., X. myosurus Mont., X. pseudoapiculata Hamme & Guerrero, X. stilbohypoxyloidea Hladki & A. I. Romero, X. telfairii, X. tucumanensis Hladki & A. I. Romero and X. xylarioides (Speg.) Hladki & Romero. From the 49 species yielded 6 cosmopolitan species, 25 pantropical species, 9 neotropical species and 9 probably endemic (YUT) species. The percentages of species according to these distributional patterns are shown in Fig. 2.

Fig. 2. Distributional Pattern (Cosmopolitan, probably Endemic from Tucuman Yungas-YUT-, Neotropical, Pantropical).

Cosmopolitan species: Astrocystis smilacicola, Kretzschmaria deusta, Poronia punctata, Rosellinia necatrix, Xylaria apiculata, X. mellissii.

Pantropical species: Biscogniauxia capnodes, Daldinia eschscholtzii, Entonaema liquescens, Hypoxylon anthochroum, H. notatum, H. rubiginosum var. microsporum, H. subrutilum, Kretzschmaria clavus, K. pavimentosa, K. sandvicense, Poronia oedipus, Rosellinia subiculata, Xylaria adscendens, X. allantoidea, X. curta, X. enteroleuca, X. fissilis, X. globosa, X. gracillima, X. grammica, X. ianthinovelutina, X. multiplex, X. myosurus, X. telfairii and X. xylarioides.

Neotropical species: Phylacia globosa, Rosellinia paraguayensis, Xylaria aff. ianthinovelutina, X. coccophora, X. cubensis, X. kretzschmarioidea, X. luxurians, X. melanura, X. pseudoapiculata.

Probably Endemic (YUT) species: Kretzschmaria argentinensis, K. sigmoidirima, Stilbohypoxylon macrosporum, S. minus, Xylaria culicicephala, X. filiformioidea, X. microceras var. yungae, X. stilbohypoxyloidea, X. tucumanensis.

Local mycobiota compared with those from other regions:

In Table 1 the species richness (n) per genus is shown comparing areas of different countries/ continents from neotropical, paleotropical and holartic kingdoms. In addition, the total number of species for the American countries and Tucuman (Argentina) is displayed through a graphic with a curve of the respective area of each country in Fig. 3. Brazil is not included because of the relation between the knowledge of the xylariaceous fungi and its large territory has not been an aim of a deeply exploration.

Fig. 3. Percentages of Hosts.

Hosts

Of all the species studied (49), two grow on dung and for 22 of them we were unable to identify the host. The identification of most of the hosts was difficult due to the advanced state of decomposition at the time of collecting. However, 23 taxa could be recognized and were 85% angiosperms with remarkable dominance of woody plants and 15% are gymnosperms (Fig 4). Classification according to The Plant List 2012 (http://www.theplantlist.org-accessed 27-07-2012):

Fig. 4. Number species/areas of the countries.

a) ANGIOSPERMS

a.1)DICOTYLEDONS

Fam. Betulaceae: Alnus acuminata Kunth (=A. jorullensis var spachii (Regel)Winckler), "aliso"

Fam. Bignoniaceae: Arrabidaea coralina (Jacq.) Sandwith, Jacaranda mimosifolia D. Don

Fam. Cannabaceae: Celtis L., Celtis tala Gillies ex Planch. (= Celtis ehrenbergiana (Klotzsch) Liebm.)

Fam. Lauraceae: Cinnamomum porphyrium (Griseb.) Kosterm. (=Phoebe porphyria (Griseb.) Mez).

Fam. Fabaceae: Anadenanthera Speg., Anadenanthera colubrina (Vell.) Brenan var. cebil (Griseb.) Altschul., "cebil colorado", Cailliea Guill. & Pert, Gleditsia L., Gleditsia triacanthos L., Mimosa L., Piptadenia Benth, Tipuana Benth.

Fam. Malvaceae: Ceiba insignis (Kunth) P. E. Gibbs & Semir (= Chorisia insignis Kunth.), "palo borracho"

Fam. Myrtaceae: Eugenia L.

Fam. Onagraceae: Jussiaea L.

Fam. Rhamnaceae: Scutia buxifolia Reissek

Fam. Rubiaceae: Psychotria carthagenensis Jacq.

Fam Rutaceae: Citrus aurantium L.

Fam. Sapindaceae: Allophylus edulis (A. St. -Hil., A. Juss. & Cambess.) Radlk.

Fam. Solanaceae: Vassobia breviflora (Sendtn.) Hunz.

Fam. Verbenaceae: Duranta L.

a.2) MONOCOTYLEDONS

Fam Arecaceae: Syagrus Mart.

Fam. Poaceae: Saccharum officinarum L.

Fam. Smilacaceae: Smilax campestris Griseb., "zarzaparrilla blanca"

b) GYMNOSPERMS

Fam. Pinaceae: Pinus taeda L.

Fam Podocarpaceae: Podocarpus parlatorei

Pilg., "pino del cerro"

The relationship between each species from Tucuman Yungas (Argentina) and their respective host species is shown in Table 2.

Table 2
Xylariaceous species and their respective hosts

Discussion

Composition of xylariaceous mycobiota

Eleven genera were recognized. If we take only the number of species into account, the best represented genus was Xylaria (n=27). On the contrary, Astrocystis, Biscogniauxia, Daldinia, Phylacia and Entonamea were the genera represented by only one species. Nevertheless it is worthwhile mentioning that only five species of the genus Xylaria are probably endemic while Kretzschmaria (n=6) and Stilbohypoxylon (n=2) have higher percentages of endemism (33.33% and 100 % respectively).

Most of the genera with a known tropical and subtropical distribution (Rogers 2000; Stadler et al. 2010; Laessee et al. 1989) are present in the studied area except for Camillea Fr. and Thamnomyces Ehrenb. It should be noted that Camillea sulcata (Starb.) Lloyd is found in the subtropical region of the Northeast of Argentina (Laessee et al. 1989).

Among the taxa with a pantropical distribution, Poronia oedipus has a wide distribution in Argentina (Buenos Aires, Catamarca, Entre Ríos, Santiago del Estero and Tucumán provinces). However, there is no record of this fungus from the southern Cone of South America. It is probably due to the lack of expeditions in the neotropical region. Recently, Agüero et al. (2011) identified P. punctata (L.) Fr. in Catamarca province. This species is categorized as vulnerable by IUCN (http://www.iuncnredlist.org/soedfiles/poronia-punctata.pdf).

Annulohypoxylon and Nemania species are generally recorded with a worldwide distribution (Rogers 2000; Lodge 2001). However, species of these two genera were not found among the nine taxa classified with this kind of distribution in the Argentine explored area.

From all the genera listed in Table 1, Phylacia Lév. is the only one which could be limited to the neotropics (Rodrigues & Samuels 1989).

Kretzschmaria is a typical genus from tropical and subtropical areas. This can be demonstrated through the number of species in the neo-and paleotropical regions, in contrast to the low diversity in the temperate European region.

The 18% of the species could be classified as probably endemic for the phytographical region Argentine "Las Yungas", which represents one of the regional ecosystems of the mayor relevance because of its high diversity of vegetal resources (Brown 1995). This value is relatively high in comparison, for example with a Neotropical zone much explored as Mexico have a 6.1 % of endemic xylariaceous species (Lodge 2001).

Comparison of the local mycobiota with others regions

As it can be expected when the xylariaceous mycobiotas from different spots (Table 1) is confronted, the neotropical kingdom is where the members of this family reach the highest diversity on Earth. This fact has been pointed out by several workers (Rogers 2000).

Vasilyeva et al. (2012) pointed out two tropical centers of biodiversity of pyrenomycetous fungi can be identified. The Western Hemisphere is the region in and around the Caribbean Sea and the Gulf of Mexico, whereas in the Eastern Hemisphere occurs in Southeast Asia.

The Western Hemisphere encompasses the neotropical region which shows the major diversity of xylariaceous fungi even in its southernmost part, as it is the area objective of the present study.

In Fig 4, it can be observed that the countries close to the center of biodiversity Caribbean Sea and the Gulf of Mexico like Panamá, Cuba, Ecuador, Venezuela and México have a larger surface than Tucumán. However the diversity of species does not increase proportionally in relation to their size. For example, Panama triples in surface Tucuman but the relation between the studied species do not increase in the same proportion (49 species in Tucumán vs 75 species in Panama).

The high diversity in Tucumán could indicate the existence of a centre of diversity of Xylariaceae in South America. Although the Brazilian data seem not to support this hypothesis, it is necessary to say that the exhaustive studies on this family have began in recent years (Pereira et al. 2010)

These data should be cautiously interpreted, because the site covered in each study substantially differs, for example, while mycobiotic studies in Venezuela (Dennis, 1970; Rogers et al., 1988) and Mexico (Pérez-Silva 1972, 1975; Medel et al. 1999, 2008, 2010; San Martín & Lavin, 1997; San Martin & Rogers 1989, 1993, 1995) cover large areas, researches in Sulawesi (Rogers et al. 1987), Papua New Guinea (Van der Gucht 1995, 1996) and Tucumán are limited to small areas, although heavily sampled. Similar situation occurs in Uganda (Taligoola &

Whalley 1976, 1977, Whalley et al. 1988) where the research was restricted to hypoxiloid species of forest areas. Zaire has a low biodiversity due probably to the fact that Dennis (1961, 1963) based his studies on specimens donated by other researchers who collected probably larger and more colorful stromata.

In Brazil main researches were concentrated in the south-Minas Gerais, Paraná, Rio Grande do Sul, Santa Catarina, Säo Paulo states, where many collections were made and the list of species present was presented by Poroca (1986) and Meijer (2006).

Hosts

Among the dicots, Fabaceae is remarkable as a host for xylariaceous species because six of its genera (Anadenanthera, Cailliea, Gleditsia, Mimosa, Piptadenia, Tipuana) are colonized by Hypoxylon subrutilum and Xylaria enteroleuca. H. subrutilum seems to be host specific with members of Fabaceae, as it was recorded by Ju & Rogers (1996), in Bolivia occurring on wood of Mimosa spp. and in Hawaii on Acacia koa A. Gray; on the contrary, X. enteroleuca seems not be host specific as it has been registered on Macadamia sp. -Fam. Proteaceae-by Callan & Rogers (1990), on Gleditschia triacanthos -Fam. Fabaceae-by Spegazzini (1898) and on Betula sp. -Fam. Betulaceae-by Martin (1970).

Cinnamomum porphyrium (Lauraceae), an endemic (Argentina-Bolivia) species, was a host colonized by 8 different species (Daldinia eschscholtzii, Kretzschmaria clavus, Xylaria globosa, X. aff. ianthinovelutina, X. fissilis, X. melanura, X. mellissii and X. telfairii). Members of this family are not common hosts for xylariaceous species, the previous record is Hypoxylon anthochroum growing on Ocotea veraguensis (Meisn.) Mez (Carmona et al. 2009), Daldinia macaronesica in Ocotea foetens and D. palmensis in Laurus azorica (Stadler et al. 2004c); and Hypoxylon canariense in Erica arborea (Stadler et al. 2008).

In Gymnosperms: Podocarpus parlatorei (endemic in Argentina-South Bolivia) was colonized by more species (7: Daldinia eschscholzii, Kretzschmaria deusta, Rosellinia subiculata, X. grammica, X. mellissii and X. pseudoapiculata) than Pinus L. (2: X. curta and X. telfairii). Among these species R. subiculata (Petrini 2003), X. curta and X telfairii (Hladki & Romero 2007) were the only ones which previously registered on gymnosperms. This amount of species (7) on a Gymnosperm is remarkable taking into account that teleomorphs of Xylariaceae occur very rarely on this group of plants in temperate areas where they are however dominating.

We identified three families of monocots: Arecaceae with Kretzschmaria clavus, Poaceae with Rosellinia paraguayensis and Smilacaceae which was colonized by Astrocystis smilacicola (= R. smilacina) and Xylaria xylarioides. Our results agree with those provided by San Martin et al. (1998 ) who pointed out most of the tropical species of Rosellinia tend to grow mainly on monocots. Another coincidence is the relation between Xylaria species and monocots as they are known to be endophytes of monocots (Petrini & Petrini 2005). On the other hand, K. clavus has a large range of hosts.

San Martin & Rogers (1995) studied the host relationship with Mexican species of the genera Xylaria, proved that most of the host were dicots (76%). Our results agree with those data from Mexico. They did not find any species growing on gymnosperms, although X. curta, X. grammica, X. mellissii (as X. arbuscula) and X. telfairii are present in the Mexican mycobiota but growing on dicots.

K. clavus is the etiological agent of the rot root of Macadamia being one important disease in Hawai (Ko et al. 1977). Hladki & Romero (2001) have recorded it for Argentina (Buenos Aires, Misiones and Tucuman) where Cinnamomum porphyrium and Anadenanthera colubrina (Vell.) Brenan are attacked by this species.

K. deusta is a pathogen on woody angiosperms from temperate zones of the Northern hemisphere (Rogers & Ju 1998), but in Argentina it was found on a native gymnosperm (Podocarpus parlatorei) and on an unidentified Myrtaceae. In addition, it was observed decline and death of plants of Citrus paradisi Macfad in a commercial farm in Salta, Argentina by K. deusta (Palacio et al. 2008).

Xylaria mellissii and X. curta were the two species that colonized highest number of hosts, wood of angiosperms and gymnosperms.

The Xylariaceae family is very well represented in this small area studied in Argentina where fourty nine species belonging to the genera Astrocystis, Biscogniauxia, Daldinia, Entonaema, Hypoxylon, Kretzschmaria, Phylacia, Poronia, Rosellinia, Stilbohypoxylon and Xylaria were recorded. In comparison with other similar neotropical regions, La Yungas shows to have a xylariaceous mycobiota with high diversity. Twenty-three host taxa are registered, mostly of them were dicot but two native trees, Cinnamomum porphyrium and Podocarpus parlatorei, and with an endemic distribution (Argentina-Bolivia) were the hosts more colonized.

Acknowledgments

We thank the directors of herbaria BAFC, BPI, C, FH, GZU, ICN, IMI, K, LIL, LPS, NY, PC, S, W, WSP for kindly providing material. The authors wish to thank Marc Stadler and Jacques Fournier for the critical revision of the manuscript. We express our appreciation to the authorities of Fundación Miguel Lillo, Tucumán, for financial support. This is publication N° 191 of the PRHIDEB-PROPLAME, partially financed by the Argentine National Research Council.

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Original recibido el 14 de agosto de 2012;
primera decisión: 15 de octubre de 2012;
aceptado el 10 de noviembre de 2012.

Editor responsable: Carlos Urcelay.