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Home , Antarctomyces, Arthrobotrys, Ascobolaceae, Ascobolus, Ascobolus carbonarius, Ascodesmis

2 : , Orbiliomycetes

1 DONALD H. PFISTER

CONTENTS 5. ...... 47 6. Glaziellaceae...... 47 I. Introduction ...... 35 7. ...... 47 II. Orbiliomycetes: An Overview...... 37 8. Karstenellaceae...... 47 III. Occurrence and Distribution ...... 37 9. ...... 47 A. Trapping 10. ...... 48 and Other Invertebrates...... 38 11. ...... 48 B. Saprobic Species ...... 38 12. ...... 49 IV. Morphological Features ...... 38 13. ...... 49 A. Ascomata ...... 38 14. ...... 49 B. Asci...... 39 15. ...... 49 C. Ascospores ...... 39 XIII. Growth in Culture ...... 50 D. Paraphyses...... 39 XIV. Conclusion ...... 50 E. Septal Structures ...... 40 References...... 50 F. Nuclear Division ...... 40 G. Anamorphic States ...... 40 V. Reproduction ...... 41 VI. History of Classification and Current I. Introduction Hypotheses...... 41 VII. Growth in Culture ...... 41 VIII. Pezizomycetes: An Overview...... 41 Members of two classes, Orbiliomycetes and IX. Occurrence and Distribution ...... 41 Pezizomycetes, of Pezizomycotina are consis- A. Parasitic Species ...... 42 tently shown in molecular phylogenetic studies B. Mycorrhizal Species ...... 42 to diverge at the base of the Pezizomycotina C. Saprobic Species ...... 42 X. Morphological Features ...... 42 phylogeny (Gernandt et al. 2001; Hansen and A. Ascomata ...... 42 Pfister 2006; Spatafora et al. 2006). Kumar et al. B. Asci...... 43 (2012) give ultrastructural data on the septal C. Ascospores ...... 43 structure, wall construction, and nuclear D. Paraphyses...... 43 division that suggest the Orbiliomycetes repre- E. Septal Structures ...... 44 F. Anamorphic States...... 44 sent the earliest diverging lineage of the Pezi- XI. Reproduction ...... 44 zomycotina. The relationship of these two XII. History of Classification and Current classes and the family diversity within the Pezi- Hypotheses...... 44 zomycetes are shown in Fig. 2.1. These two A. Families of the Pezizomycetes ...... 46 groups show very few shared characteristics 1. ...... 46 2. Ascodesmidiaceae...... 46 other than the formation, for the most part, of 3. ...... 46 well-developed apothecial ascomata in which 4. ...... 46 asci are generally arranged in a hymenium with paraphyses. Orbiliomycetes was recog- nized as a class only recently (Baral 2003). 1Farlow Herbarium and Library of Cryptogamic Botany, Department of Organismic and Evolutionary There is a single order, Orbiliales, and a single Biology, Harvard University, Cambridge, MA, 02138, USA; family, , with perhaps 300 species e-mail: dpfi[email protected] currently arranged in three teleomorphic

Systematics and Evolution, 2nd Edition The Mycota VII Part B D.J. McLaughlin and J.W. Spatafora (Eds.) © Springer-Verlag Berlin Heidelberg 2015 36 D.H. Pfister

Fig. 2.1 A diagrammatic representation of the relation- received maximum parsimony, posterior probability, ships of the Pezizomycetes and Orbiliomycetes, includ- and maximum-likelihood support 75 %, 95 %, and ing the families of the Pezizomycetes, based on Hansen 75 %, respectively, are in bold et al. (2013) and Pfister et al. (2013). Branches that Pezizomycotina: Pezizomycetes Orbiliomycetes 37 genera. An early diverging position of the class In this chapter, the two classes are treated has been confirmed by morphological studies independently in separate sections. and molecular phylogenetic analyses (Kumar et al. 2012; Spatafora 2006; Wang et al. 2006). Previously, the Orbiliaceae was placed among II. Orbiliomycetes: An Overview the as a family based on Nannfelt’s (1932) classic work on the nonlichenized inop- The family Orbiliaceae was proposed by Nann- erculate . The Pezizomycetes feldt (1932) and was restricted to those fungi have long been recognized to include a single with small so-called inoperculate asci, minute order, the , and include 16 families ascospores, and a waxy texture. The asci and and approximately 1,200 species. The group, paraphyses are often agglutinated. Nannfeldt in whole or in part, has been the subject of illustrated several species showing details of recent surveys (Hansen and Pfister 2006; Har- the thin-walled globose to angular cells of the rington et al. 1999; Perry et al. 2007; Pfister excipulum. A monograph on the et al. 2008). by Svrcek (1954) included 16 species that were In regard to morphological characteristics distinguished based on ascospore size and and ecology, the two classes also are distinct. shape, the color of the ascomata, and the form The Orbiliomycetes are saprobic and in some of the paraphyses. The groundbreaking work of cases supplement their nutrition through the Hans-Otto Baral and his collaborator, Guy Mar- capture and consumption of small inverte- son, has led to a refinement of the family and brates. Their are small, nonseptate recognition of many more species. Kirk et al. (save for a single example), uninucleate, and (2008) proposed 288 species, but the extent of contain a membrane-bound structure, the the diversity to be described is in flux. A mono- body (Baral 1994; Benny et al. 1978; graph has been in preparation by Baral for Kumar et al. 2012). Spores are discharged many years. Modern methods employ morpho- actively. Asci are small and generally lack a logical and histochemical characters previously well-developed apical apparatus. Anamorphic not used and incorporate ecological characters states are frequently formed, and their number as well. Of particular note is the reliance on has increased steadily in the literature in recent fresh, living material (Baral 1992). There has years. Species are found on dung, on wet wood been a burst of activity focusing on the descrip- on the forest floor, in water-soaked areas where tion of teleomorphic taxa and associated ana- they are somewhat ephemeral, or on dead wood morphs (e.g., Kohlmeyer et al. 1998; Liu et al. hanging in trees where, because of their 2005a, b, 2006; Mo et al. 2005; Pfister 1994; drought resistance, they are persistent (Baral Pfister and Liftik 1995; Qiao et al. 2011; Qin 2011). No species are plant parasitic. et al. 2010; Su et al. 2011; Tanabe et al. 1999; The Pezizomycetes are saprobic, plant par- Webster et al. 1998; Wu et al. 2007; Yang and asitic, or associated with plants as ectotrophic Liu 2005; Yu et al. 2007a, b, c, 2011; Zhang et al. mycorrhizae (Hansen and Pfister 2006; Teder- 2007), and in this work there has been an soo et al. 2006; Wei et al. 2010). The ascomata intense interest in the diversity and functional are always to some degree ephemeral. Spores morphology of the anamorphic states. This is are large, generally more than 10 mm and may particularly the case regarding those that reach a length of 60 mm, and single-celled. capture nematodes, i.e., species referred to Spores generally are actively discharged and allied form genera. through a well-developed opening created by the rupture of the ascus wall forming an oper- culum that is either terminal or eccentric in position. The trufflelike habit has developed III. Occurrence and Distribution independently in several lineages across the class (Læssøe and Hansen 2007). These hypo- Members of the Orbiliomycetes are found geous taxa generally do not have active spore around the world in boreal to tropical habitats. release. Furthermore, several species complexes, such 38 D.H. Pfister as O. luteorubella and O. xanthostigma, that addition to trapping nematodes, some hypho- represent morphologically very similar collec- mycetes in the orbiliaceous lineage trap other tions seem to be found broadly distributed small invertebrates such as collembolans and around the world and across latitudes. There rhizopods (e.g., Barron 1981; Drechsler 1936, are two major ecological groups—those that 1937b). Even though these fungi may occur on occur in moist habitats such as on dung, old wood or cellulose-rich substrates, there is evi- polypores, and moist wood on the forest floor dence that few of the -trapping fungi or near streams and species that are drought are able to break down cellulose (Park et al. tolerant. These drought-tolerant species are 2002). found on dead twigs and small woody branches in the canopy of living trees. Often species of both ecological types are found in association B. Saprobic Species with unicellular green algae, but there is no evidence that the fungi are lichenized. Until the work of Baral (2011) and Marson, most studies of diversity within the Orbiliomycetes were focused on those species on moist sub- A. Species Trapping Nematodes strates. The discovery that these fungi are and Other Invertebrates highly diverse in the dry habitats of hanging branches in living trees has greatly expanded Since the classic work of Drechsler (1937a)on the number of species that have been described the nematode-trapping , these through intense collecting in such habitats. fungi have attracted many researchers to this Although these species are treated here as fascinating system. The work of Barron (1977, saprobic, there is little clear evidence of their 1981) further set the stage for ecological and mode of nutrition. Drechsler (1938) discussed descriptive work on these fungi. Pfister (1994, at length hyphomycetes that are parasitic on 1997) and Pfister and Liftik (1995) documented oospores of Oomycota; these also seem now to the life history connection between these be clearly referable to the Orbiliaceae. These nematode-trapping hyphomycetes and teleo- fungi were “considered from a morphological morphic members of the Orbiliomycetes. The viewpoint... [to] fit acceptably in the preda- several anamorphic form genera involved are cious series” (Drechsler 1938). Whether they discussed in what follows. fall in the main clade of nematode trapping Trapping of nematodes and other inverte- Orbiliaceae or outside it remains to be studied, brates is accomplished by the formation of but that a variety of life styles and biotic modes constricting rings, hyphal networks, or sticky are found within the family is unquestionable. knobs. These trapping structures are often, but It is important to note as well that no plant not exclusively, initiated in the presence of parasitic species have been discovered so far prey. When nematodes become ensnarled in in this lineage. the trap, the then grows and penetrates the body of its prey and proliferates within it. Conidia are formed in the presence or absence IV. Morphological Features of nematodes and are generally dispersed across the medium, that is, they are not con- A. Ascomata centrated around trapped nematodes (Barron 1977). Earlier classification schemes for these Apothecial ascomata range in size from 0.5 to fungi relied primarily on conidial morphology, 5 mm and are saucer-shaped to pulvinate. particularly spore shape and septation. Phylo- Macroscopically they have been described as genetic studies have indicated that the form of appearing waxy at sight or to touch. This is the trapping devices gives important informa- a reference at least in part to their shiny, trans- tion for classification (Hagedorn and Scholler lucent appearance. The sterile tissues of the 1999; Rubner 1996; Scholler et al. 1999). In ascomata are composed of an outer layer, or Pezizomycotina: Pezizomycetes Orbiliomycetes 39 ectal excipulum, of globose or angular cells that enhanced magnification. Spores are globose, in some species give rise to superficial hairs. ellipsoidal, ovoid, reniform, acicular, or filiform These cells may contain one or more cytoplas- and mostly smooth, although spore wall mic bodies that may take on several different ornamentations are known, for example, in shapes but disappear when material is mounted O. xanthostigma. Ascospores of the Orbiliomy- in KOH. Cells in the outer excipulum may be cetes contain an inclusion, the spore body. This encrusted with granules or produce glassy pro- membrane-bound body was suggested to derive jections or the cells may be embedded in an from the plasma membrane by Baral (2003), amorphous matrix. The inner layers of the but Kumar et al. (2012) and Benny et al. ascomata are of interwoven hyphae or com- (1978) showed that the spore body is derived pressed angular cells (Nannfeldt 1932). from mitochondria. The spore body is attached to the cell membrane and is surrounded by a rough endoplasmic reticulum (Benny et al. B. Asci 1978). The contents are electron dense (Benny et al. 1978). Under a light microscope it is seen The asci of members of this class are small cylin- in unstained living spores as a refractive body. drical to broad cylindrical with or without cro- Its appearance can be enhanced by application ziers; in some cases they have elongate, forked of aqueous brilliant cresyl blue or other vital bases. Asci contain eight to many ascospores per stains. It disappears in dead spores and can be ascus. Asci are inamyloid. The apex ranges from destroyed by application of KOH. The spore hemispherical conical to truncate. Walls are thin, body is a synapomorphy for the class and can composed of two layers in transmission electron take on many different forms, from globose to microscopy (Benny et al. 1978; Kumar et al. tear-shaped to elongate. Its shape and orienta- 2012). The apex is generally little developed. tion is consistent within the ascospores of a Benny et al. (1978) reported no pores; Kumar particular species. The shape and position of et al. (2012) found an electron-light zone at the the spore body is considered a critical taxo- apex. In some taxa, an opening develops on the nomic character. Generally spores have a single shoulder of the truncate asci (Zhang et al. 2007), spore body, but an example of a spore with two and this was confirmed by Kumar et al. (2012). In spore bodies is known (Wu et al. 2007). As some species, the apical walls may be thickened mentioned previously, its contents are electron to form a cap (Kohlmeyer et al. 1998), as in dense; it does not appear to be composed of Orbilia subgenus Hemiorbilia,inwhichthe lipids but may be proteinaceous (Kumar et al. apex of the ascus is thickened (Baral 2011). In 2012). No function has been attributed to the all cases, asci appear to dehisce by irregular tear- spore body, but Benny et al. (1978) offered ing. Kumar et al. (2012)foundsimilarities three possible functions: for flotation and bal- between the asci of the Orbilia species they stud- ance in a liquid environment before and during ied and members of the . Asci germination, storage of waste products, and and paraphyses may be embedded in a gelati- storage of materials for use during germination. nous matrix that causes them to stick together None of these functions has been documented, and not fan out when crushed for microscopic and ultimately the function of the spore body observation. This is particularly the case in mem- may have to do with special metabolic activities bers of the genus Hyalorbilia. within the spore.

C. Ascospores D. Paraphyses

The initial striking feature of the ascospores is Paraphyses are sparingly septate, broad, some- their small size. They range in size from 3 to times with a prominent capitate terminal cell. 8 mm. Observation of them is difficult in some They may be encrusted or contain refractive situations and requires oil immersion and materials. In some species, the paraphyses and 40 D.H. Pfister asci are closely adherent and are difficult to and the other adhesive traps, but, importantly, separate. these fungi form a single clade. Yang et al. (2007) proposed a scheme for the evolution of trapping structures. E. Septal Structures Other form genera are those that produce staurosporous or helicosporous conidia or are Ascus, ascogenous, and nonascogenous hyphae associated with aquatic habitats. These include have simple septa, with septal pores plugged by Angulospora (Pfister 1997;Webster1992; unelaborated electron-dense, nonmembranous Webster and Descals 1979), (reports occlusions. Globose Woronin bodies were summarized by Pfister 1997), located on both sides of the septum (Kumar (Kohlmeyer et al. 1998), Helicoon (Pfister 1997), et al. 2012). Trinacrium (Matsushima 1995), and Triden- taria. Others are found on a variety of plant materials in terrestrial habitats, and these F. Nuclear Division include (Chen et al. 2007a, c), Dac- tylellina (Li et al. 2005; Qin et al. 2010), Division is characterized by the retention of an (Chen et al. 2007a, b, c; Pfister 1997; Qin et al. intact nuclear envelope and a two-layered disk- 2010), and Pseudotripoconidium (Yu et al. 2011). shaped spindle pole body (Kumar et al. 2012). Staurosporous conidia and those reported from other aquatic anamorphs are detected in stem flow, through-fall, and treeholes (Go¨nczo¨l G. Anamorphic States and Re´vay 2003, 2004, 2006;Re´vay and Go¨nczo¨l Many members of the Orbiliomycetes have 2011). Among these are conidia referred to been shown to produce mitospores in culture; form genera of anamorphs associated with they are hyphomycetous or synnematous. All species of Orbiliomycetes. These include spe- conidia are formed by holoblastic wall exten- cies of Angulospora, Dactylaria, Dicranidion, sion. A variety of named form genera have been Dwayaangam, Trinacrium, and Tridentaria. applied to these conidial states. Anamorphs can Of these Go¨nczo¨l and Re´vay (2004) found provide some useful information on the biology that Trinacrium in particular was prevalent and ecology of these fungi. Those that have and that Dwayaangam and Trinacrium species been demonstrated to trap nematodes and were particularly diverse and geographically other invertebrates fall into the following widespread (Go¨nczo¨l and Re´vay 2006). Such genera: Arthrobotrys (Li et al. 2005; Mo et al. observations suggest that canopies are impor- 2005; Pfister 1994, 1997; Pfister and Liftik 1995; tant habitats for these fungi. We expect that Qiao et al. 2011), Drechslerella (Li et al. 2005), these are likely Orbilia anamorphs that are Dwayaangam (Barron 1991), , associated with the drought-tolerant, canopy- known only in its anamorphic state (Scholler inhabiting Orbiliomycetes, which are fungi on et al. 1999), Lecophagus (Tanabe et al. 1999), dead branches in many woody plants. Go¨nczo¨l and (Li et al. 2005; Pfister and Re´vay (2006) say, regarding their findings 1997). There remains debate about the applica- of aquatic hyphomycetes in these terrestrial tion of some of these names. For general habitats, that “this reinforces our belief that overviews the interested reader may consult an ecological group of hyphomycetes distinct Hagedorn and Scholler (1999), Liou and Tzean from those in lotic habitats exists and functions (1997), and Scholler et al. (1999). Through a in canopies.” By extension, we might assume multigene analysis Yang et al. (2007) postulated that branches and twigs inhabited by Orbi- that two lineages evolved trapping mechanisms, liomycetes are a major part of this canopy with one lineage giving rise to constricting rings ecosystem. Pezizomycotina: Pezizomycetes Orbiliomycetes 41

V. Reproduction ascus, the ectal excipulum, spore bodies, the presence or absence of gelatinous material in Although many of the Orbiliomycetes have the hymenium, and pigmentation [see Zhang been grown in culture, few reports have been et al. (2007) for a summary of the genera]. published on apothecial production in the lab. Spontaneous production of ascomata has been noted (Drechsler 1937a; Rubner 1994). Zachar- VII. Growth in Culture iah (1983) was able to induce ascomatal primorida. Breeding systems have not been One of the most remarkable features of the studied in part because of the unpredictability Orbiliomycetes is the ease with which most of of producing ascomata in culture. the species can be cultivated on standard media. Spores deposited on the surface of agar media germinate readily, with the exception of VI. History of Classification and the species of Hyalorbilia for which there are Current Hypotheses only a few reports of successful cultivation. Spores germinate quickly, and growth is often luxuriant on substrates containing glucose. The family Orbiliaceae was created in 1932 Sporulation is common under room condi- in Nannfeldt’s (1932) classic treatment of the tions. inoperculate discomycetes. Recognizing the special features of this group, he removed the genera Orbilia and Hyalina from the core group VIII. Pezizomycetes: An Overview of inoperculate discomycetes. Primary treat- ments of the family were those by Svrcek (1954) and Spooner (1987). Nannfeldt’s cir- The Pezizomycetes comprise a single order, cumscription of the family was used with few Pezizales, with 16 families currently recognized. changes until the 1990s. The placement of the The full diversity of the order has not yet been family outside the lineage initi- completely recognized in the classification, and ally began to be questioned by Pfister (1997), no doubt other lineages will of necessity be and subsequent work has confirmed the place- named. Ascomata are both epigeous and hypo- ment as an early diverging lineage within the geous. The epigeous types are apothecial, cleis- Pezizomycotina (Gernandt et al. 2001; Spata- tothecial, or highly reduced, being composed of fora et al. 2006). The position of the Orbiliomy- only a few asci in clusters on vegetative hyphae cetes as an early diverging lineage in the with little or no excipular tissue. In epigeous filamentous along with the Pezizo- lineages, spores are generally forcibly dis- mycetes was one of the major surprises in the charged with the asci rupturing by the forma- era of molecular phylogenetic studies. tion of an operculum. Hypogeous members Within the Orbiliomycetes there is a single occur in most families containing mycorrhizal order, and only three teleomorphic genera are members. To date, more than 15 independent recognized. Other genera may well be proposed origins of trufflelike members are known as more of these fungi come to be known within a majority of the families (Læssøe and through wider collection of them and collec- Hansen 2007). tions come under deeper scrutiny. The described genera are Orbilia, the largest and most diverse genus with two subgenera and IX. Occurrence and Distribution nine sections in Baral’s treatment (2011), Hya- lorbilia with six to eight species (Baral and Pezizomycetes can be found around the Marson 2000), and Pseudorbilia, with a single world, but representatives are unevenly species (Zhang et al. 2007). These genera are distributed. Members of certain families, such distinguished based on characteristics of the as the Pezizaceae, Morchellaceae, Helvellaceae, 42 D.H. Pfister

Rhizinaceae, and many of the Pyronemataceae, generally do not form rhizomorphs (Tedersoo show a particularly high diversity in tempera- et al. 2006; Wei et al. 2010). Mycorrhizal species ture regions. Others are more abundant in seem to predominate in areas that experience tropical areas, as exemplified by the Sarcoso- xeric conditions (Smith et al. 2006). Mycorrhi- mataceae and Sarcoscyphaceae. Recent studies zal lineages are often also those that contain have made clear that, although collecting in this truffle or trufflelike taxa. There are no truffle- group is robust in many areas of the Northern like species in the families Sarcoscyphaceae and Hemisphere, the Southern Hemisphere is Sarcosomataceae, and likewise there are no poorly documented. Nutritionally, Pezizomy- known mycorrhizal species. cetes are saprobic or mutualistic (Tedersoo et al. 2006) or are parasitic on bryophytes and vascular plants (Hansen and Pfister 2006). C. Saprobic Species Many of the species collected on soil occur in disturbed areas or in soil that has a high pH and Pezizomycetes occur on organic material of low content of organic matter. Petersen (1985) various types—decaying wood, dung, leaf litter, provides a review of the edaphic factors and twigs. The diversity of Pezizales on dung is involved in growth and reproduction. well documented, with many studies and keys available (Bell 1983; Doveri 2004; Richardson and Watling 1997). Some clades are nearly A. Parasitic Species exclusively found on dung, most notably spe- cies of Ascobolaceae (Brummelen 1967). Dung- The plant parasitic species are scattered in var- inhabiting species are found in several clades of ious lineages. undulata is a serious the Pyronemataceae and Pezizaceae (Hansen root parasite of (Ginns 1968). and Pfister 2006; Hansen et al. 2001; Perry species may cause dieback on conifers. Calos- et al. 2007). In most cases, little is known of cypha fulgens is implicated as a seed pathogen the biology of saprobic species. - of conifers (Paden et al. 1978; Salt 1974). ceae, Sarcosomataceae, and Chorioactidaceae craterium has been implicated as the causal are found exclusively on wood, leaves, and agent of canker in (Davidson plant debris. 1950). Phymatothrichopsis ominivora is a root pathogen of cotton and other dicotyledonous plants and is known only in its anamorphic X. Morphological Features state (Uppalapati et al. 2010). A clade in the Pyronemataceae, including and A. Ascomata , is parasitic on bryophytes (Han- sen and Pfister 2006). Apothecia occur on the Apothecia are the basic type of ascomata. These thalli or surrounding soil if they are rhizoidal range in size from several millimeters to several parasites (Do¨bbler 1979). centimeters and vary from sessile cups of small to large size to stalked cups to the stipitate piliate structures found in the Helvellaceae, B. Mycorrhizal Species Discinaceae, and Morchellaceae. Some highly reduced members are little more than small With the advent of molecular approaches to fascicules of asci such as in . Further studying mycorrhizal root tips a number of reductions include highly reduced forms found examples of mycorrhizal Pezizomycetes have in Eleutherascus and Monascella (Guarro and been documented, and it is now known that Arx 1986; Stchigel et al. 2001). Here the asci are mycorrhizal lineages are found throughout the formed in clusters on unspecialized hyphae. class (Tedersoo et al. 2006). Morphologically, Ascomata are often highly pigmented, particu- the mycorrhizae have a mantle that is often larly the hymenium. Carotenoid pigments have thin, have a well-developed Hartig net, and been characterized (Arpin 1969). Pezizomycotina: Pezizomycetes Orbiliomycetes 43

Variations are found in typical apothecial in the form of an operculum, and its position, ascomata. On the one hand, there are highly the reaction of the wall layers in various stain- reduced types that are merely asci scattered ing agents in light microscopy, and the ultra- on hyphae as mentioned previously, but small structure of the ascus wall are all important cleistothecial ascomata have also been found in characters. Amyloid asci are found in - a few cases (Hansen et al. 2005b). In the case of ceae and Ascobolaceae (Hansen et al. 2001), but Orbicula parietina, active discharge is lost and this character has been lost several times, most spores are presented in a powdery mass. Hey- notably in the hypogeous members of the Pezi- denia species form stalked cleistothecial fruit zaceae. Amyloid asci are not found outside bodies closely related to O. parietina (Lecht- these families. mann and Cle´menc¸on 2011). The ascomata of The layering of the wall as seen in transmis- Heydenia, which was recently placed in the sion electron microscopy helps delimit groups, Pyronemataceae (Lechtmann and Cle´menc¸on particularly the taxa with thick lateral walls in 2011), deserve special comment. H. alpina is a the Sarcoscyphaceae and Sarcosomataceae. The small fungus that occurs on plant debris and asci of these fungi were termed suboperculate . It is stipitate with a dark that at by Le Gal (1946a, b, 1953). Subsequently, the top expands to hold a pale mass of hyaline Ecblad (1968) and Samuelson (1975) dis- spores and radiating hyphae. No asci are pres- counted the term. ent, but molecular evidence confirms that this Not all members of the class have opercula. species belongs in the Pyronemataceae as a Hypogeous (Læssøe and Hansen 2007) and close neighbor of Orbicula, another fungus some cleistothecial taxa (Hansen et al. 2005a, that produces dry powdery masses of spores b) have lost the operculum and spores remain at maturity. The two genera share a cleistothe- either within asci until eaten and the fruit body cial fruit body and asci that seem to disintegrate is broken down, or the asci disintegrate and a early in the development of the ascomata but powdery mass is formed. diverge on important morphological features (Lechtmann and Cle´menc¸on 2011). These genera, often placed outside of the Pezizales, C. Ascospores have been demonstrated through molecular systematic studies to be members of the order. No member of the Pezizomycetes has septate Truffleoid fruit bodies are found in several spores at maturity. Septa may form in germinat- lineages. Following the terminology of Weber ingspores,butonlywhenagermtubehas et al. (1997), these are stereothecia, exothecia, already been established. The spores may take or ptycothecia, depending on the arrangement on a variety of shapes from globose to naviculate, of the asci and the presence or absence of with smooth or ornamented surfaces. Generally, well-developed hymenial layers. Truffle-type the ornamentations are derived from secondary ascomata are derived from apothecial forms. wall material, although a few examples are Læssøe and Hansen (2007) summarize much known of spore ornamentation without a contri- of the literature on these taxa. bution of secondary walls. The spore walls are Some highly reduced forms are found multilayered and may be thin or thick (Merkus among bryophyte parasites. The fruit bodies 1976). The number of nuclei per spore varies are partly closed and have only a few asci. from one, two, or four to many, and such varia- They give the appearance of perithecia. tion has been shown to have some taxonomic value (Berthet 1964;Korf1972, 1973).

B. Asci D. Paraphyses Brummelen (1978, 1994) summarized the structural characters of the ascus in the Pezizo- The pigmentation of the hymenium is attribu- mycetes. The apical apparatus, when present, is ted to the pigments found in the paraphyses. 44 D.H. Pfister

Yellow, red, and orange carotenoid pigments Attempts to germinate mitospores from these are distinctive features of some of these fungi. mats have failed, and the function of these These pigments are dissolved in oil droplets spores remains unknown. Several anamorphic in the paraphyses. A variety of carotenoid pig- states have been discovered through molecular ments are known to occur in these fungi, as phylogenetic studies to belong within the summarized by Arpin (1969). Paraphyses are Pezizomycetes, but they have no known generally septate and may be either uninucleate teleomorphs. Most notable among these is Phy- or multinucleate, this latter character being matotrichopsis omnivora, the cotton root path- used in the classification (Berthet 1964; Pfister ogen. This falls within the Rhizinaceae based on et al. 2008). In some cases, paraphyses are inter- molecular studies, but despite intensive study spersed with hyaline or darkly pigmented, often of this important pathogen, no evidence of a thick-walled elements or setae. Examples of teleomorph has been discovered. Another such setae are found in members of the Rhizi- example is Cephaliophora, a dung-inhabiting naceae, Sarcosomataceae, Chorioactidaceae, fungus and one that has been implicated in and Sarcoscyphaceae. causing keratitis (Hoog 2000).

E. Septal Structures XI. Reproduction

Ultrastructural characters of septal pore plugs Extensive studies have been carried out on sev- and Woronin bodies correlate to some degree eral members of the class. was early with families and lineages in the order. Kim- incorporated as a genetic tool. has brough (1994) summarizes much of the infor- also been extensively studied. Early cytological mation. Septal pores in vegetative hyphae studies on members of the Pyronemataceae generally have lamellae embedded in a matrix. were used to explore mitotic events in asci. These septa generally have associated Woronin bodies. The Woronin bodies take on several forms—globular, hexagonal, or cylindrical, sometimes considerably elongate. The form of XII. History of Classification and the Woronin bodies is an important taxonomic Current Hypotheses and phylogenetic character. The septa at the base of the asci also provide important charac- The current classification has its origins in the ters. These septa are occluded by dome-shaped, work of Boudier (1885, 1907), who was the first pyramidal, or dumbbell-shaped structures. to use the presence of an operculum to define These plugging structures show electron-dense this group. Hypogeous, truffloid members pre- bands, lamellae, and raylike extensions. viously treated in the order Tuberales have now been placed in the Pezizales based on morphol- ogy, cytology (Trappe 1975), and molecular F. Anamorphic States sequence analyses (Læssøe and Hansen 2007). Wide-ranging phylogenetic work within the Anamorphic states have been reported across order led to the recognition of three primary many families. Conidia are blastic in develop- lineages (Landvik et al. 1997). Subsequent work ment and are generally hyphomycetous. has expanded this framework and has sug- Table 2.1 provides a summary of the known gested groupings of families that in part reflect anamorphs. In many of these anamorphs, the those previously based on morphological, cyto- germination of conidia has not been observed, logical, and pigment data and are noted in suggesting that they may rather act as sperma- earlier work (Korf 1972, 1973; Rifai 1968). The tia. The investigations by Healy et al. (2013) most recent comprehensive overview of the identify anamorphic spore mats in several class is that of Hansen and Pfister (2006). Cur- clades in the Pezizaceae and Tuberaceae. rently, 15 families are recognized (Fig. 2.1), Table 2.1 Summary of known anamorphs of Pezizomycetes

Family Anamorphic form genus Teleomorphic genus References Ascobolaceae Rhizostilbella Ascobolus Seifert (1985) Papulospora Ascobolus Brummelen (1967)

An oidial state Ascobolus Brummelen (1967) 45 Orbiliomycetes Pezizomycetes Pezizomycotina: Caloscyphaceae Geniculodendron Paden et al. (1978) Chorioactidaceae Kumanasamuha geaster geaster Nagao et al. (2009) Verticicladium acicola Paden (1972) Morchellaceae Costantinella Paden (1972) Pezizaceae Glischroderma Pezizaceae Norman and Egger (1999), Hansen et al. (2001), Healy et al. (2013) Chromelosporium Peziza, , Muciturbo Hennebert (1973), Paden (1972), Warcup and Talbot (1989), Healy et al. (2013) Oedocephalum, including highly Peziza, , , Hennebert (1973), Hennebert and Belleme`re (1979), Paden (1972), reduced forms Cleistoiodophanus Bezerra and Kimbrough (1976) Pyronemataceae Actinosporella megalspora lechithina Descals et al. (1998) Alciphila vulgaria Harmaja (2002a, b) Ascorhizoctonia Yang and Korf (1985), Barrera and Romero (2001) Cephaliophora None known Tanabe et al. (1999) Complexipes Yang and Korf (1985) Dichobotrys Hennebert (1973) Micronematobotrys None known Sun and Guo (2010) “Nodulosporium”-like majalis Paden (1972) Rhizinaceae Phymatotrichopsis None known Uppalapati et al. (2010) Sarcosomataceae , , Hughes (1958), Paden (1972) latahensis Sarcoscyphaceae Molliardiomyces , Sarcoscypha, Paden (1984), Pfister (1973b) Tuberaceae Unnamed sympodulosporous Urban et al. (2004), Healy et al. (2013) type 46 D.H. Pfister though the relationship among those families in are formed on a much-reduced excipulum many instances is not well resolved. Although (Brummelen 1981). These species are saprobic this group is reasonably well known, future on dung and often are isolated from soil. Asci sampling will certainly change our understand- are inamyloid and are saccate or pyriform. ing of the relationships and of these lineages. A Ascospores are brownish and have reticulate particular focus is on the large and heteroge- spore ornamentations. Septa in ascogenous neous family Pyronemataceae, in which several cells and the ascus bases are dome-shaped lineages are recognizable based on both with arrays of radiating tubular elements and sequence analyses and expanded knowledge of a striate structure associated with the septal nutritional modes, morphology, and anamor- pore rim in vegetative cells (Brummelen 1989; phic states. Kimbrough 1994). Four genera are placed in the General overviews of the families can be family (Hansen et al. 2005b). A recent study by found in Cannon and Kirk (2007). The follow- Hansen et al. (2013) places this family within ing augmented descriptions include available the larger and highly diverse sister group to the structural and cytological information that has Pyronemataceae sensu stricto. been used in refining the classification.

3. Caloscyphaceae A. Families of the Pezizomycetes The brightly colored cupulate ascomata stain blue green when bruised. No hypogeous mem- 1. Ascobolaceae bers are known. Caloscypha fulgens is a parasite Only epigeous members are known, and these of seeds and seedlings. The ascomata occur primarily on dung, but a few species are have a well-developed inner layer of interwoven found on plant material; hyphae and an outer layer of globose or angular is found on burned material (Brummelen cells. Ascospores are globose, and their walls 1967). All species are saprobic and ascomata are smooth (Harmaja 2002a). The genus Calos- have multiple cell layers. Developmental pat- cypha was previously placed in the Pyronema- terns have been characterized by Brummelen taceae, in part because of its carotenoid (1967). Cleistothecial developmental types are pigments and because of the globose spores found. Asci blue diffusely along the walls in (Arpin 1969; Korf 1972, 1973). For details on iodine solutions. Ascospores are uninucleate septal construction see Kimbrough and Curry (Berthet 1964); in Ascobolus and (1986b). Recently a second genus was added to the spore walls are ornamented with purple the family, Kallistoskypha, with a single species, brown pigments that become fissured at matu- K. incarnata, which is associated with Eucalyp- rity (Brummelen 1967). The placement here of tus found around the Mediterranean region species, with their hyaline ascos- (Pfister et al. 2013). pores, has been suggested on morphological grounds and has been confirmed in molecular phylogentic studies (Landvik et al. 1997). Septa 4. Chorioactidaceae associated with the ascogenous system are The ascomata are cupulate or deep urnulate dome-shaped (Kimbrough 1994; Kimbrough and dark on the outer surface; the hymenium and Curry 1985). Three genera are generally is beige, brownish, or orange. No hypogeous recognized. members are known in this family, and all the species are considered to be saprobic. The asco- 2. Ascodesmidiaceae mata have a well-developed inner layer and an outer layer of globose or angular cells that give Ascomata are much reduced in most genera; for rise to thick, brown hairs encrusted with gran- example, in Eleutherascus species the asci are ular material. The asci are inamyloid and thick- scattered on hyphae. In Ascodesmis, several asci walled with a terminal operculum. Ascospore Pezizomycotina: Pezizomycetes Orbiliomycetes 47 ornamention is in the form of warts or ribs. et al. 2006). The ascomata are characteristically Four genera have been included (Pfister et al. constructed with an inner layer of interwoven 2008). hyphae and an outer layer of elongate cells oriented perpendicularly to the outer surface. Asci are inamyloid. Ascospores are tetranucle- 5. Discinaceae ate (Berthet 1964), smooth, or ornamented with The ascomata of this family are discoid or gyro- irregular warts, and Woronin bodies are elon- mitroid. Both epigeous and hypogeous mem- gate, hexagonal, or globose (Kimbrough 1994). bers are known. Many species in the family are Septa at the base of the asci are electron- mycorrhizal (Tedersoo et al. 2006, 2010). The opaque. There are hemispherical structures ascomata are characteristically constructed that become cone- to dumbbell-shaped with with an inner layer of interwoven hyphae and V-shaped striations; in vegetative cells, an an outer layer of elongate cells oriented perpen- electron-translucent torus separates the pore dicularly to the outer surface. Asci are inamy- plug from the septal pore border. Vegetative loid, and ascospores are tetranucleate (Berthet cells possess globular or slightly angled Woro- 1964), smooth, or elaborately ornamented with nin bodies (Kimbrough 1994; Kimbrough and warts, often with apiculae. Woronin bodies are Gibson 1989; Kimbrough et al. 1996). elongate and often surround the septal pore. The septa at the base of asci are accompanied by an electron-opaque, hemispherical structure 8. Karstenellaceae that becomes cone- to dumbbell-shaped with In this family, the ascomata are little more than V-shaped structures; an electron-translucent a very thin, resupinate crust situated on leaf torus separates the pore plug from the septal litter and wood surfaces. Asci are scattered on pore border (Kimbrough 1991, 1994). The this mycelial mat. Nothing is known of its mode septal ultrastructure is similar to that of the of nutrition. Asci are inamyloid and operculate. Helvellaceae. Generally, five or six genera are Ascospores are binucleate or multiguttulate recognized. (Hansen et al. 2008; Harmaja 1969). No ultra- structural details are known for the vegetative septal construction or for the septa of the asci. 6. Glaziellaceae Molecular data place it in an unresolved posi- The ascomata in this family are large and hol- tion near Helvellaceae and Tuberaceae. There is low with a basal opening. The unisporic, clavate a single genus with a single species recognized, to globose asci are embedded in the rindlike Karstenella vernalis. wall. Ascus walls break down, but the spores remain embedded in this peridiumlike wall. The single species, aurantiaca, is pre- 9. Morchellaceae sumed to be mycorrhizal. See Gibson et al. The morels are characterized by stipitate (1986) for ultrastructural details that initially apothecia, with an interrupted hymenial layer supported the placement of Glaziella in the giving a honeycomb appearance, or by cupulate Ascomycota. forms such as Disciotus. Hypogeous members include Leucoangium (Li 1997) and Kalapuya 7. Helvellaceae (Trappe et al. 2010). Species are saprobic and have been implicated in mycorrhizal relation- Ascomata are sessile cupulate or stipitate cupu- ships (Buscot 1994; Dahlstrom et al. 2000). Asci late, saddle-shaped, or columnar (Abbott and are inamyloid. Ascospores are multinucleate Currah 1997; Dissing 1966). Both epigeous and (Berthet 1964) and lack both prominent wall hypogeous species are found in the family ornamentation and internal oil droplets. Tradi- (Hansen and Pfister 2006; Kimbrough et al. tional classifications of the species accept six to 1996). Many or most are mycorrhizal (Tedersoo eight species, but phylogenetic studies indicate 48 D.H. Pfister the presence of many species (Du et al. 2012; phyletic and will need to be broken down into Kuo et al. 2012; Taskin et al. 2010) that seem several additional genera (Hansen et al. 2001, restricted to specific continents (O’Donnell 2005a, b). et al. 2011). Some of these species are morpho- logically distinct; others are cryptic (O’Donnell et al. 1997, 2011). The septal construction is 11. Pyronemataceae similar to that found in the Helvellaceae. At the base of the asci, dome-shaped structures Pyronemataceae is the largest of the families of with V-shaped striations are found at the septal the class and the most diverse ecologically and pore. Lamellate structures are found within morphologically. As recognized here, the family septal pores of most vegetative cells. Elongate includes Aleuriaceae, Humariaceae, Otideaceae, Woronin bodies have been found along with and Geneaceae. These fungi are saprobic, hexagonal ones (Kimbrough 1994). Five or six mycorrhizal vascular plants or parasitic on genera are recognized. bryophytes. Epigeous members are cupulate, often possessing hairs on the outer surface. Hypogeous members are scattered in the family. 10. Pezizaceae The ascomata show considerable variation in construction and development. Asci are inamy- Both epigeous and hypogeous taxa are found; loid, and ascospores vary in shape and orna- there are multiple origins of the hypogeous taxa mentation from smooth to variously within several clades (Hansen et al. 2001, 2002, ornamented (Perry et al. 2007). Several different 2005a; Healy et al. 2009). Saprobic members septal types are present; at least four types of include those on dung, wood, and other plant septa plugging have been associated with asci debris. Mycorrhizal taxa are found in several (Kimbrough 1994). These are the aleurioid type, lineages. Some taxa are found exclusively on which have a granular, opaque matrix that bor- burned areas. The ascomatal structure varies, ders both the ascal and ascogenous hyphal sides but multiple layers composed of large, thin- of pores; later this becomes fan-shaped with a walled globose cells are often present. Asci gen- lamellate electron-translucent torus adjacent to erally become blue in iodine solutions. This the pore rim. The otideoid type, is characterized bluing can take several forms. The reaction by double translucent bands in a granular, may be diffuse along the entire wall or concen- opaque pore matrix. Later the pore plug is dif- trated in the upper portion of the ascus but ferentiated into two zones, an inner dense zone most intensely at the tip, or in the upper por- and an outer less opaque zone. The pulvinuloid tion with a more intense ring surrounding the type, is similar to the Helvellaceae at maturity opercular region (Hansen et al. 2001). The blu- with V-shaped striations, but these are fewer ing reaction has been lost in several taxa, and less prominent than in Helvellaceae. The including some that are hypogeous. Ascospores scutellinioid type has a large hemispherical sep- are uninucleate (Berthet 1964), with or without tal pore plug that becomes zonate; the inner secondary wall ornamentations (Hansen et al. zone adjacent to the pore lumen is electron- 2001, 2002). Oil droplets may or may not be opaque, and an outer, thicker zone is composed present in the ascospores. Septa in vegetative of less dense material. In general, in this family, cells are lamellate with large globose Woronin vegetative cells have a lamellate structure in the bodies. At the ascus bases, septa are occluded septal pores. Woronin bodies are small and by convex or biconvex bands that become cov- largely globular, but often hexagonal. ered with electron-opaque amorphous material The pyronema type has asci and ascogen- or by an additional secondary wall (Curry and ous hyphae that are similar to those of the Kimbrough 1983; Kimbrough 1994; Kimbrough Ascobolaceae but with a different electron den- et al. 1991). Approximately 30 genera are cur- sity core and small, radiating tubular bands rently recognized in the family, but it is well similar to those of Ascodesmis (Kimbrough established that the genus Peziza is nonmono- 1994; Kimbrough and Curry 1986a, b). Pezizomycotina: Pezizomycetes Orbiliomycetes 49

The family includes approximately 80 Li and Kimbrough 1995b). Twelve genera are genera. It is perhaps highly heterogeneous and commonly recognized within this family. one that will see future division and refinement. Romero et al. (2012) review the classification The family was recently studied by Hansen et al. within the family. (2013), and this work provides the most accu- rate overview of the family. 14. Sarcosomataceae

12. Rhizinaceae In this family, ascomata are cupulate or urnu- late, tough and leathery of moderate to large The ascomata in members of this family are flat, size, and black or dark brown on the outside, recurved, or pulvinate. is a often with a paler hymenium. No hypogeous pathogen of conifer roots. Also included here is members are known. Most species are pre- , whose species are found on water- sumed to be saprobic on plant debris, but in soaked wood and which is presumed to be a the case of Urnula, the species is said to be saprobe (Pfister 1973a). The construction of the parasitic. The ascomata have a well-developed ascomata follows a pattern of interwoven inner layer, with gelatinous material, and an hyphae forming the medulla, with more densely outer layer of globose or angular cells that give interwoven hyphae contributing to the cortical rise to dark hyphoid hairs. Asci are inamyloid layer. These fungi are characterized by indeter- and thick-walled with a terminal operculum. minate marginal growth. The asci are inamy- The ultrastructure of the asci and ascospores loid, and ascospores are smooth or ornamented have been studied by Belleme`re et al. (1990). with warts and apiculae. The spores are pre- The ascospores are smooth or with wrinkled or sumed to be tetranucleate. Studies are needed folded walls or with low warts. In addition, they on the septal structure. The family encom- are multinucleate. Septal structures are com- passes the two genera mentioned earlier as posed of an electron-dense matrix and globose well as Phymatotrichopsis omninvora, which is Woronin bodies that are often accompanied by known only as an anamorph. short or long, cylindrical, hexagonal Woronin bodies. Septa at the ascus bases are composed of a dumbbell-shaped matrix with V-shaped 13. Sarcoscyphaceae bands. The septal structure was studied by Li and Kimbrough (1995b). Five or six genera are Ascomata are generally cupulate, bright col- commonly accepted. ored, stipitate, or sessile. No hypogeous mem- bers are known; all are considered saprobic. The ascomata have a well-developed inner 15. Tuberaceae layer and an outer layer of globose or angular cells or of tightly interwoven hyphoid cells ori- Until recently, all members of this family were ented parallel to the outer surface. Cells on the considered to be hypogeous. The recent place- outer surface sometimes give rise to single or ment of Nothojafnea thaxteri among these fasciculate hairs. Ascospores are smooth or fungi is notable as the first epigeous member ornamented with ridges and ribs. In vegetative of the family (Bonito et al. 2013). The truffles cells, lamellate structures are poorly differen- of commerce ( and tiated or missing. Septa in vegetative cells are T. magnatum) belong here. Ascomata are imperforate, plugged by an electron-opaque, highly convoluted and folded, often with cham- fan-shaped matrix surrounded by a number of bers or pockets of asci. The large, primarily globose electron-dense Woronin bodies. The Northern Hemisphere genus Tuber has been Woronin bodies are globose or occasionally investigated by Bonito et al. (2010). All mem- hexagonal and electron-opaque. At maturity bers are mycorrhizal on a range of plant obscure V-shaped striations may be found families (Tedersoo et al. 2006). The asci are (Benny and Samuelson 1980; Kimbrough 1994; inamyloid, globose, or pyriform with one to 50 D.H. Pfister eight spores per ascus. The ascospores, in addi- behavior, whereas some members of the Orbi- tion to being often multinucleate, are often liomycetes trap and consume invertebrates. elaborately ornamented with reticula, ridges, Recent studies have placed many hypogeous and warts. Septal structure was studied by Li Pezizomycetes within families. These studies and Kimbrough (1995b), who determined that highlight the evolutionary patterns within the there was some heterogeneity among those spe- class in which suites of changes, including loss cies examined. In that study, the researchers of active spore discharge mechanisms and found long, cylindrical Woronin bodies in extended ascomata developmental times, take some of the species, similar to those of the place in distantly related clades. Most hypoge- Morchellaceae, Discinaceae, and Helvellaceae, ous Pezizomycetes are found within clades with and rectangular ones in another, similar to other mycorrhizal members. The diversity in those of the Pyronemataceae. Septa associated both Orbiliomycetes and the Pezizomycetes with ascogenous hyphae and ascus bases also has proven to be greater than previously antici- differed among the sampled taxa. O’Donnell pated, and much researches remains to be done et al. (1997) included a series of genera in addi- regarding the biogeography and evolutionary tion to Tuber. Seven genera are commonly histories of the group. recognized. Bonito et al. (2013) estimate a late origin of the genus Tuber and discuss the Southern Hemisphere diversity. References

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