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Institute of Systematic Botany, Friedrich Schiller University Jena, Philosophenweg 16, 07743 Jena, Germany
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTS AND EVALUATIONDISCUSSIONLITERATURE CITED |
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The fruit body and in particular the gleba development of several species of Lycoperdaceae has been examined by light microscopic analysis of microtome sections of fruit body primordia in different ontogenetic stages. The gleba of Langer-mannia gigantea develops in a unique and previously unknown manner: the hymenium-forming palisade structures are borne on fan-like branches of hyphae. Hence the term of flabelloid glebal development is introduced. The other genera of Lycoperdaceae, including the genus Calvatia, are characterized by both lacunar and coralloid development of the gleba. Because these features cannot be distinguished clearly, this type of glebal development is referred to as coralloid-lacunar. Due to the peculiar ontogeny I suggest keeping the genus Langermannia separate from Calvatia.
Key words: fruit body development, gleba development, Langermannia gigantea, Lycoperdaceae
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND EVALUATIONDISCUSSIONLITERATURE CITED |
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, Kreisel 1962, 1967, Demoulin 1971), the developmental characters of their basidiocarps and in particular their glebal characters have been examined only rarely. The work on the ontogeny of this family published several decades ago (Rehsteiner 1892, Rabinowitsch 1894, Cunningham 1926, Lander 1933, Swartz 1933, 1935, 1936a, , Swartz b, Brandza and Solacolu 1937, Swoboda 1937, 1938, Ritchie 1948, Ahmad 1950, Marchant 1969) did not include the genus Langermannia Rostk., only the peridial features were analyzed once (Swoboda 1940). Considering the systematic relationships with the relatively well studied genus Calvatia Fr. (Swartz 1933, 1935, Brandza and Solacolu 1937), a similar type of fruit body development had been assumed.
Langermannia consists of three species (Kirk et al 2001), of which only L. gigantea (Batsch : Pers.) Rostk., the type species, is common in the temperate zone (Kreisel 1994). This species is commonly known as giant puffball. It forms one of the world’s largest basidiocarps from single primordia, a development known as monocentric (Clémençon 1997). Like all members of the Lycoperdaceae, it is a saprobiotic organism. Its fruit bodies are common on fertile meadows or in open woodland.
The exoperidium of the genus consists of only one layer; it lacks the pseudoparenchymatous endostratum typical of all other Lycoperdaceae. The dehiscence of the peridium is irregular; there is no defined ostiolum. The capillitium of the Lycoperdon-type is septated. Langermannia shares the last two features with Calvatia.
The genus Langermannia was erected by Rostkovius (1844) for species with irregular dehiscence of the peridia, allowing the mature glebal mass with its strongly developed capillitium to tumble away with the wind. He included five species into the genus, L. candida (=Calvatia candida), L. aculeata, L. flavescens and L. punctata (all = Calvatia excipuliformis) and the type species L. gigantea. When Calvatia was erected by Fries (1849) he included only the type species, Calvatia craniiformis (Schwein.)Fr. He transferred Langermannia into Lycoperdon. With the emendation of Morgan (1890) all species of Lycoperdaceae with an irregular dehiscence of the peridium, including Langermannia, were incorporated into Calvatia. Many authors followed this generic concept (Hollós 1904, Lloyd 1905, Fischer 1933, 
marda 1958, Zeller and Smith 1964). Kreisel (1962) accepted Langermannia due to the anatomy of its peridium but discussed its close relationship to Calvatia. In later publications he included it into Calvatia because of the type of peridial dehiscence and the features of the capillitium (Kreisel 1992, 1994). Many authors follow his notion (Pegler et al 1995, Winterhoff 2000) while others accept Langer-mannia as a genus (Calonge and Martín 1990, Lange 1993, Calonge 1998, Kirk et al 2001). The genus Calvatia is a nomen conservandum against Langer-mannia (Greuter et al 1999).
The systematic relationships of Langermannia and Calvatia are discussed controversially (Kreisel 1992, 1994, Calonge and Martín 1990, Lange 1993, Calonge 1998). Recent molecular systematic studies (Krüger et al 2001, Moncalvo et al 2002, Krüger and Kreisel 2003, Bates 2004, Lebel et al 2004, Vellinga 2004) could not clarify this matter but revealed close relationships of the Lycoperdaceae and the Agaricaceae. The present study contributes a novel argument for the separation of Langermannia from Calvatia by means of comparative anatomical investigation of the ontogenetic stages of several species of Lycoperdaceae.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND EVALUATIONDISCUSSIONLITERATURE CITED |
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) and stained with toluidine blue O (0.1% w/v in H2O) and safranine O (0.5% w/v in H2O). Cover slips were attached with Merckoglas (Merck). The slides were examined with a Jenalumar microscope; microphotographs were taken with a Olympus Camedia digital camera (C3040ZOOM) and evaluated with software analySIS (version 3.2). Materials.— – Bovista plumbea Pers. : Pers. : Jena (Thuringia): on a meadow near Möbis, 1.5 km south of Münchenroda, 9 Sep 2004, leg./det.: M. Gube; Bovista aestivalis (Bonord.) Demoulin: Jena (Thuringia): on foliage litter of Quercus petraea at Tännicht, 0.5 km southeast of Jenaprießnitz, 1 Oct 2004, leg./det.: M. Gube; Calvatia excipuliformis (Pers.) Perdeck: Marienberg (Saxony): on needle litter of Picea abies and between Avenella flexuosa at the edge of a path between Ansprung and Rübenau, 3 km south of Rübenau, 3 Sep 2005, leg./det.: M. Gube; Langermannia gigantea (Batsch : Pers.) Rostk.: Jena (Thuringia): beneath Quercus robur at the edge of a meadow near Möbis, 1.5 km south of Münchenroda, 9 Sep 2004, leg./det.: M. Gube; Lycoperdon foetidum Bonord.: Zittau (Saxony): on needle litter of Picea abies on the northern slope of the Lindeberg, 2.5 km northwest of Hainewalde, 3 Jul 2004, leg./det.: M. Gube; Lycoperdon lambinonii Demoulin: Lobenstein (Thuringia): on needle litter of Picea abies at the Tännig, 2 km southeast of Lobenstein, 25 Sep 2004, leg./det.: M. Gube; Lycoperdon perlatum Pers. : Pers. : Zittau (Saxony): On litter of Picea abies and Sorbus aucuparia at the northern slope of the Buchberg, 1 km southwest of Jonsdorf, 2 Nov 2003, leg./ det.: M. Gube ; Jena (Thuringia): on foliage litter of Quercus petraea and Fagus sylvatica at the northern slope of the Hirschberg, 1 km southeast of Jenaprießnitz, 1 Oct 2004, leg./det.: M. Gube; Lycoperdon pyriforme Schaeff. : Pers. : Jena (Thuringia): on a decayed stump of Fagus sylvatica inside the Langer Grund, 3.5 km west of Jena, 18 Nov 2003 and 30 Sep 2004, leg./det.: M. Gube; Vascellum pratense (Pers. : Pers.) Kreisel: Ellrich (Thuringia): on a meadow, 1 km west of Sülzhayn, 18 Oct 2003, leg./det.: M. Gube
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RESULTS AND EVALUATION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND EVALUATIONDISCUSSIONLITERATURE CITED |
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The differentiation process of the primordial plectenchyme starts in primordia of 1 mm diam. Radially oriented, frequently septated hyphae appear on the surface of the primordium to form the exoperidium (FIG. 1). The first internal differentiation occurs in primordia of 2 mm diam, where an area of gaps emerges in the central primordium, later forming the glebal cavities. Such a formation of gleba is lacunar because the gaps emerge independently inside the previously undifferentiated primordial plectenchyma, according to Fischer (1933). They arise from separation and splitting, as well as from autolysis of some hyphae. This has been discussed controversially by other authors (Rehsteiner 1892, Cunningham 1926, Fischer 1933, Lander 1933, Swartz 1933, Marchant 1969). Cavity formation extends centrifugally. In primordia of 5 x 4 mm, the gleba differentiates further as hyphae originating from the surrounding plectenchyma grow into the cavities to form a palisade layer of hyphal tips (FIGS. 2–3), shown also by Swartz (1933) and Ritchie (1948). Because this occurs centrifugally as well the central part of some cavities can be covered with palisades while the peripheral part remains undifferentiated (FIGS. 2–4). Lohwag (1925) described such a glebal formation as coralloid because this outwardly progressing differentiation leads to coral-like branches of the trama while the cavities between them usually are connected.
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). They show a gradual transition into their glebal counterparts. Therefore they can be kept apart only by their position in young stages, as already mentioned by Rehsteiner (1892).
In primordia of 9 x 5 mm and larger the centrifugally progressing differentiation reaches the endoperidial layer, which has differentiated in the meantime. The periclinally arranged hyphae of the endoperidium do not differentiate further. Therefore the peripheral part of the cavities in this region is bordered by endoperidial hyphae without palisades (FIG. 6).
The palisade layer surrounding the glebal cavities consists of clavate hyphal tips with deep staining plasmatic content (FIG. 7). These probasidia pass basally into a subhymenium, while their tips form an even surface. Therefore it is obvious that this layer forms a true hymenium later on. Yet it is referred to as palisade layer or prehymenium because no ripe basidia with spores were observed. In more mature primordia only the apical part of the palisades stains deeply due to vacuole formation while the plasma retreats toward the apical region (FIG. 8). This is typical for the senescence of hymenial palisades (Marchant 1969). The subhymenial plectenchyme remains thin-walled and equally frequent septated during differentiation (FIGS. 7–8). In the subgleba it is weakly developed and is compressed due to growth of the cavities (FIG. 9). Further inside the trama branches hyphae are arranged in parallel bundles (FIGS. 7–9). Their cell walls are thicker, they stain metachromatic and usually are septated sparsely. In the gleba some of these hyphae form the latter capillitium while they develop into the walls of the persistent chambers in the subgleba. The hyphae of the endoperidium are differentiated similarly; there are even transitions where trama branches meet it (FIGS. 6).
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). In mature primordia however the subgleba differs from the gleba; because the palisade layer shows other signs of senescence they are lacking plasmatic content then and become thick-walled (FIG. 9). In addition the cavities stretch along the boundary region of gleba and subgleba in later stages.
Features of the glebal development of Bovista, Calvatia, and Vascellum deviating from Lycoperdon.— –
Species of Bovista, Calvatia and Vascellum have similar glebal ontogeny compared to Lycoperdon. Species of Bovista differ somewhat in the features of the palisades; they are larger and more capitate than those of Lycoperdon, Vascellum or Calvatia (FIG. 10). Primordia of Calvatia excipuliformis develop distinctly slower than those of other Lycoperdaceae. With primordia at 4 mm diam, when the palisades usually emerge, only the first signs of cavities are noticeable. Fully developed palisades may not be found in fruit bodies smaller than 25 x 12 mm.
Peculiar features are the distinct diaphragm in mature basidiomata of Vascellum and the pseudo-diaphragm in some species of Calvatia, both separating gleba and subgleba (Kreisel 1962). These structures could not be observed in the examined specimens. However the cavities of more mature basidiomata of V. pratense (7 x 6 mm) and C. excipuliformis (25 x 12 mm) are stretched along the border of the already distinguishable glebal and subglebal plectenchyme (FIGS. 11–12). Diaphragm and pseudodiaphragm arise from stretched glebal and subglebal cavities. This was observed by Rabinowitsch (1894), whereas Cunningham (1926) supposed these structures to be formed from subglebal cavities only.
Gleba development of Langermannia gigantea.— –
L. gigantea shows a gleba formation quite different from that of the other Lycoperdaceae. In the earliest examined stages (at 7 mm diam) the palisade layer does not cover the surface of cavities but was found to be carried on fan-like branched hyphae (FIG. 13). For them the term of flabelloid structures is introduced here. They form a nearly continuous, irregularly lobed zone with the palisade layer growing outward into the undifferentiated plectenchyme (FIGS. 14) and sideways, thus forming secondary cavities at later stages (FIGS. 15–16). In primordia (at 11 mm diam) more palisade layers emerge inside while the primordium grows. The palisade layers replace only undifferentiated plectenchyme. Hyphae transmitting the palisade structures often are observed (FIG. 17). These are probably remains of the replaced plectenchyma because they are more common in the periphery, where palisades are newly formed. While the basidiocarp grows the outer zone of palisades extends and the most exterior parts of the undifferentiated plectenchyme are stretched to form the peridia. The differentiation into exo- and endoperidium was not yet detectable. Also the definition of boundaries between glebal and peridial plectenchyma could not be observed at the stages examined.
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). Several of these originate from one multibranched hypha, which is itself a branch of higher order (FIG. 13). Therefore a whole fan may be traced back to a few or even a single subhymenial hypha. Therefore great areas of the glebal trama show a low density of hyphae (FIGS. 15–16). Only in the surrounding of the secondary cavities do hyphae exist resembling the central trama hyphae of other Lycoperdaceae (FIG. 15). They are relatively thin-walled but stain metachromatic and are more sparsely septated, in contrast to the subhymenial hyphae. The compact subgleba, which is weakly developed even in mature fruit bodies, cannot be observed in the primordia.
Visibility of dolipore septa.— –
At the septa of subhymenial and central tramal plectenchyme of most Lycoperdaceae, as well as in the undifferentiated plectenchyme of Langermannia gigantea, large dolipore complexes up to 2 µm diam are visible (FIGS. 7–8, 17). This also can be observed in primordia of Agaricus bisporus (Lge.) Sing., indicating that this staining method is not unique for Lycoperdaceae (FIG. 18). The high visual quality of the dolipore structures surpasses that of most other staining methods. The exceptional size and stainability of the dolipore complexes might have been caused by the aldehyde fixation, which can induce swelling of dolipore complexes (Hoch and Howard 1981).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND EVALUATIONDISCUSSIONLITERATURE CITED |
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) or lacunar type (Fischer 1933). This system of classifying gleba development of gasteromycetes has been criticized because of numerous intermediates between the extremes (Reijnders 1976, 2000). However it still is widely accepted (Kreisel 1969, Dring 1973, Jülich 1981, Miller and Miller 1988). Coralloid gleba development is considered to be a typical character for the Lycoperdaceae (Kreisel 1962).
Gleba development of the genera Lycoperdon, Bovista, Vascellum and also Calvatia however does show elements of both the coralloid and the lacunar type. On the one hand many cavities arise without connection to other already existing cavities, particularly in the beginning of gleba formation and in the subgleba, implying lacunar development. On the other hand the differentiation processing outward and the connections between several cavities point toward coralloid development. In addition the examined species show neither a continuous, coral-like branched glebal cavity, as illustrated by Fischer (1933) for coralloid development, nor completely isolated chambers, as described by Lohwag (1925) for the lacunar type.
These features have been described in most publications on the development of Lycoperdaceae (Rehsteiner 1892, Rabinowitsch 1894, Cunningham 1926, Lander 1933, Swartz 1933, 1935, 1936a, , Swartz b, Brandza and Solacolu 1937, Swoboda 1938, Ritchie 1948, Ahmad 1950, Marchant 1969, Reijnders 1976). Gleba development was interpreted in many different ways, ranging from a generally coralloid (Kreisel 1962, Fischer 1936) to predominantly lacunar (Ahmad 1950) and everything in between (e.g. Fischer 1933). Neither coralloid nor lacunar correctly describe this development, where features of both types obviously merge. Therefore the coralloid-lacunar type of gleba development is introduced here for the Lycoperdaceae.
The gleba formation of Langermannia gigantea deviates from hitherto known types. There are no primary glebal cavities, but the palisades themselves equal those of other Lycoperdaceae. The flabelloid structures bearing the palisades may form secondary cavities or remain as palisade zones among undifferentiated plectenchyme. Such a zone is formed in the region of the gleba facing the later peridia. This type of gleba development is referred to as flabelloid.
Yet another type of gleba development has been proposed for Calvatia bicolor (Lév.) Kreisel (Swoboda 1937). In this species the gleba is described as being composed of coralloid, thin branches of hyphal bundles bearing basidia directly attached to them and thus not forming a true hymenium. Since then C. bicolor has not been analyzed to verify Swoboda’s observations. In addition no other examined species of Lycoperdaceae show similar features, so the record remains doubtful.
In addition to the type of gleba development several other features are unique to Langermannia. It is the only genus of the Lycoperdaceae with only two-layered rhizomorphs, lacking cortex hyphae. They strikingly resemble those of Agaricus (Agerer 2002, Gube 2005) instead of the three- to four-layered rhizomorphs typical for Lycoperdaceae (Townsend 1954, Cairney et al 1988, Agerer 2002, Gube 2005). The one-layered exoperidium lacking pseudoparenchyma is unique within this family. It is present however in the Mycenastraceae (Hansen 1962, Kreisel 1962), which are supposed to be closely related (Bates 2004). Finally, even in primordia at 11 mm diam, exo-and endoperidia are indistinguishable while they are clearly differentiated in primordia of other Lycoperdaceae when reaching that size.
The features that are common to Langermannia and Calvatia are shared by other members of the family. The typical dehiscence of the peridia is also common in Vascellum pratense, although not as distinctive.
The septated capillitium of the Lycoperdon-type (Kreisel 1962) can be observed in both Langermannia and some species of Calvatia. Species of Calvatia with unseptated capillitium and slit-like pits in the capillitial threads have been separated by Kreisel (1989, 1992) forming the genus Handkea. Many authors do not accept this genus (Calonge and Martín 1990, Lange 1993, Calonge 1998, Kirk et al 2001); the slit-like pores are products of rupturing along the true, fine pores in the thin-walled capillitium, according to Lange (1993). In addition true septa occur in both genera as well as in the capillitium of species of Lycoperdon and Disciseda, although there they are found only scarcely (Kreisel 1962).
Recent work on the molecular systematics of Lycoperdaceae supports a clade of Langermannia gigantea, Calvatia pachydermica (Speg.) Kreisel and C. bicolor (Bates 2004). These species share several morphological features that differ from Calvatia s. str. (Kreisel 1992, 1994). However the fruit body development described for C. bicolor (Swoboda 1937) differs from both L. gigantea and other species of Calvatia. Other molecular systematic studies could not reveal the position of Langermannia within the Lycoperdaceae (Hibbett et al 1997, Krüger et al 2001, Moncalvo et al 2002, Krüger and Kreisel 2003, Lebel et al 2004, Vellinga 2004) because no or only few members of Calvatia or Langermannia were examined. Considering the available data Langermannia clearly differs in many important features from Calvatia and therefore should be referred to as a separate genus.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Present address: Microbial Phytopathology, Institute of Microbiology, Friedrich Schiller University Jena, Neugasse 25, 07743 Jena, Germany. E-mail: matthias.gube{at}uni-jena.de 
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