**3. Bamboo ascomycetes: history**

**Figure 1.** Ascostromata of *Shiraia bambusicola* collected from China.

mushroom and usually called as "bamboo pith." It has been cultivated worldwide and brings a high economic income. Some of bamboo fungi, however, are pathogens, most of which are

Cultivation of economically important bamboos is often threatened by fungal infection and diseases which eventually result from serious damages on bamboo cultivation [4]. A number of foliage diseases (e.g., leaf spots and leaf blight) of bamboos have been recorded. However, fungi cause comparatively less damage to bamboo than culm diseases [5]. For example, during 1988–1990, 5300 hectares of bamboos were affected by *Balansia take* (Miyake) Hara which occurred on bamboo culms in Fujian, China. More than 200,000 bamboos were cut down and burned to prevent the pathogen from spreading. However, this caused serious economic losses. In 1960, Hino and Katumoto explained that the relative importance of different diseases affecting bamboos is difficult to assess because of the general lack of information accompanying the disease records [4]. Thus, pathologists have now started paying attention on the research of bamboo pathogens. A book entitled *Diseases of bamboos in Asia: an illustrated manual* listed 122 fungal diseases in Asia region, with 100 records in India and 15 in Thailand [5]. In China, 183 fungal pathogens associated with bamboo were recorded [6, 7]. Numbers of ascomycetous fungi recorded as bamboo pathogens were quite high. However, in the absence of molecular data, most of the fungal names recognized remained artificial. A sizable number of

*Shiraia bambusicola* Henn. is one of the famous bamboo pathogens because of its medicinal value. The fungus produces large, pinkish ascostromata on living bamboo branches (**Figure 1**). Their fruiting bodies, as a traditional Chinese medicine, are used for curing rheumatoid arthritis [8], as well as extracting the metabolite, hypocrellin [9], which has promising applications

ascomycetes. In this chapter, a review of bamboo ascomycetes is provided herein.

**2. Importance of bamboo ascomycetes**

166 Bamboo - Current and Future Prospects

taxa were not even identified up to species level.

The term "fungorum bambusicolorum" (bambusicolous fungi), was first used by Iwao Hino [14], though the author did not give a definition. "Bambusicolous" means "living on bamboo" [3]. Kevin D. Hyde and colleagues in 2002 defined bambusicolous, which embodies fungi growing on any bamboo substrates, including leaves, culms, branches, sheathes, flowers, rhizomes, and roots [3]. Subsequently, the phrase "Bambusicolous fungi" has been widely used by mycologists worldwide [15, 16].

Lembosia Léveillé is the first mycologist, who mentioned the presence of a fungus on bamboo. In 1845, he described *Roumegueria goudotii* (Lév.) Sacc. ex Clem. & Shear (Basionym: *Dothidea goudotii* Lev.) occurring on leaves of *Chusquea* sp. and *Sphaeria bambusae* Lev. collected from the culms of *Bambusa bambos* (L.) Voss, in Tolima, Columbia [17]. Later, this author introduced two new species *Asterina microscopica* Lev. and *S. hypoxantha* Lev. from the leaves of *Chusquea* sp. and culms of *B. bambos*, respectively, in 1846 [18].

In 1854, Miles J. Berkeley recorded *Hypoxylon fuscopurpureum* (Schwein.) Berk. from *Phyllostachys* sp. and *Sasa* sp. [19]. The next year, Jean P.F.C. Montagne introduced a new species *S. fusariispora* Mont. on leaves of *Bambusa* sp. [20]. During 1871–1880, eight new ascomycetes were described from bamboo host, and between 1881 and 1920, the numbers got increased up to more than 100 species (**Figure 2**). However, the number of newly described species declined before and after the Second World War [3] (**Figure 2**). Nevertheless, during 1951–1990, publications of new species on bamboo got increased dramatically (**Figure 2**). Iwao Hino and Ken Katumoto made an earlier significant contribution on bambusicolous fungi during 1960–1970s, by recording 104 new species of ascomycetes [4, 21–29]. In 1961, Iwao Hino wrote *Icones fungorum bambusicolorum japonicorum*, and recorded 460 ascomycetous species worldwide, of which 175 species were from Japan [30]. In the following years, more records and checklists of fungi on bamboo were carried out by mycologists. Petrini Orlando and colleagues in 1989 mentioned 63 records from France [31]. Ove E. Eriksson and Yue Jinzhu published *Bambusicolous pyrenomycetes*, *an annotated checklist* and listed 587 ascomycetous taxa in 1998 [32]. A checklist for 104 species from China is provided in 1999 by [33].

& K.D. Hyde, *Occultabambusa* D.Q. Dai & K.D. Hyde, *Parabambusicola* Kaz. Tanaka & K. Hiray, *Pseudoastrosphaeriella* Phookamsak et al., *Pustulomyces* D.Q. Dai & K.D. Hyde, *Seriascoma* Phook., D.Q. Dai & K.D. Hyde, associated with bamboo substrates were introduced by the

A Review of Bambusicolous Ascomycetes http://dx.doi.org/10.5772/intechopen.76463 169

The morphological characters are important in fungal identifications. Most fungi produce their fruiting bodies on hyphae with two phases of reproductions (sexual and asexual reproduction). Thus, when an ascomycetous fungus bears its fruiting body by sexual reproduction, it usually produces an ascoma (**Figure 3**), and on the other hand, it produces either a conidioma or hyphomycetous fruiting structures. The major morphological characters of sexual morph are the type of ascomata, hamathecium, asci, and ascospores, and those of asexual

The ascomata of bambusicolous fungi have various types, no matter what shapes or colors. They can form on bamboo leaves, culms, or even sheathes (**Figure 4**), with the positions being immersed, erumpent, or superficial (**Figure 5**). Some ascomata are stromatic, with unilocule to multilocules. Ascomata produce asci and ascospores within, when mature. An ostiole, where ascospores release, is commonly present at the top of an ascoma. Peridium is the wall of an ascoma and usually is composed of several layers of angular cells. The hamathecium is

**4. Major morphological characters of bamboo ascomycetes**

morph are conidiomata, conidiophores, and conidiogenous cells [63, 64].

the sterile tissue, formed by hyaline filaments which are called paraphyses.

**Figure 3.** Line drawing of a fruiting body (ascoma) of sexual morph of ascomycete.

natural classification [16, 45, 46, 52, 53, 55, 57–62].

**4.1. Morphological characters of sexual morph**

*4.1.1. Ascomata*

**Figure 2.** Number of fungi described from bamboo until 2017.

Recent contributions to the knowledge of bamboo-associated ascomycetes were the following. An article in 2000 reviewed 189 species of fungi, of which were 80 ascomycetes, 40 coelomycetous, and hyphomycetous fungi on bamboo substrates [34]. Kevin D. Hyde and colleagues reviewed bamboo fungi in detail in 2002 [3]. A total of 80 fungi were recorded on submerged bamboo in 2003, see [35].

During 2003–2008, the Japanese mycologists carried out a series of studies on bambusicolous fungi in Japan and introduced around 25 new taxa based on the morphological characters [15, 36–42]. In 2007, authors analyzed molecular diversity of bamboo-associated fungi from Japan, based on 257 endophytes strains, isolated from bamboo tissues; however, most isolates were not identified to species level [43]. Until 2009, the significant phylogenetic analysis was first used to classify the new taxa by [44].

Currently, during 2011–2017, approximate 145 new species and new records belonging to 65 genera, 37 families have so far been described or reported by mycologists [16, 45–52] based on taxonomic and phylogenetic analyses. In the short period of 6 years, six new families *viz. Anteagloniaceae* K.D. Hyde & Mapook [53], *Bambusicolaceae* D.Q. Dai & K.D. Hyde [54], *Occultabambusaceae* D.Q. Dai & K.D. Hyde [3], *Parabambusicolaceae* Kaz. Tanaka & K. Hiray [52], *Pseudoastrosphaeriellaceae* Phookamsak & K.D. Hyde [55], *Roussoellaceae* J.K. Liu et al. [56]. and 21 new genera viz. *Amphibambusa* D.Q. Dai & K.D. Hyde, *Bambusaria* Jaklitsch et al., *Bambusicola* D.Q. Dai & K.D. Hyde, *Bambusistroma* D.Q. Dai & K.D. Hyde, *Botryobambusa* Phook., J.K. Liu & K.D. Hyde, *Brunneoclavispora* Phook. & K.D. Hyde, *Embryonispora* G.Z. Zhao, *Flammeascoma* Phook. & K.D. Hyde, *Gregarithecium* Kaz. Tanaka & K. Hiray, *Kalmusibambusa* Phookamsak, Tennakoon, *Multilocularia* Phook., Ariyaw. & K.D. Hyde, *Neoanthostomella* D.Q. Dai & K.D. Hyde, *Neogaeumannomyces* D.Q. Dai & K.D. Hyde, *Neoramichloridium* Phookamsak et al., *Neoophiosphaerella* Kaz. Tanaka & K. Hiray., *Neoroussoella* J.K. Liu, Phook. & K.D. Hyde, *Occultabambusa* D.Q. Dai & K.D. Hyde, *Parabambusicola* Kaz. Tanaka & K. Hiray, *Pseudoastrosphaeriella* Phookamsak et al., *Pustulomyces* D.Q. Dai & K.D. Hyde, *Seriascoma* Phook., D.Q. Dai & K.D. Hyde, associated with bamboo substrates were introduced by the natural classification [16, 45, 46, 52, 53, 55, 57–62].
