Ascomycota
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{{Taxobox | color = lightblue | name = Ascomycetes | image = Morchella rotunda 040410w.jpg | image_width = 238px | image_caption = Morchella rotunda | regnum = Fungi | divisio = Ascomycota | subdivision_ranks = Subphyla/Classes | subdivision =
- Archaeascomycetes Template:Ref labelTemplate:Ref label
- Euascomycetes
- Hemiascomycetes
- or
- or
- Pezizomycotina Template:Ref labelTemplate:Ref label
- Laboulbeniomycetes
- Lecanoromycetes
- Leotiomycetes
- Pezizomycetes
- Sordariomycetes
- (and many more)
- (Discomycetes)
- Saccharomycotina
- Taphrinomycotina
}}
Members of the Division Ascomycota are known as the Sac Fungi and are fungi that produce spores in a distinctive type of microscopic sporangium called an ascus (Greek for a "bag" or "wineskin"). This monophyletic grouping was formerly known as the Ascomycetae and is an extremely significant and successful group of organisms (12,000 species in 1950), accounting for some 75% of all described fungi.Template:Ref labelTemplate:Ref label Included are most of the fungi that combine with algae and sometimes cyanobacteria to form lichens. The majority of fungi that lack morphological evidence of sexual reproduction are placed here or in the Deuteromycota. Better known examples of sac fungi are yeasts, morels, truffles, and Penicillium. The majority of plant-pathogenic fungi belong to this group, or the Deuteromycota. Species of ascomycetes are also popular in the laboratory. Sordaria fimicola, Neurospora crassa and several species of yeasts are used in many genetics and cell biology experiments.
An ascomycete produces great numbers of asci at any one time, and these may be contained in a structure called an ascocarp (also called an ascoma, this is the 'fruiting body'). Each ascus usually contains eight (or a multiple of 8) ascospores, the result of one round of mitosis following meiosis. The resulting haploid nuclei are surrounded by membranes (from the plasma membrane in Euascomycetes; from the nuclear membrane in Hemiascomycetes) and eventually a spore wall.
An exception to the structure described above are ascomycetous yeasts, which are secondarily unicellular.
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Explanation of subclassification Template:Ref label
- The subphylum Pezizomycotina contains all the Ascomycota which have ascocarps ('fruiting bodies'), except for one genus, Neolecta, which is classed in the Taphrinomycotina. So this group includes almost all the macroscopic ascos. The cells reproduce by fission, not by budding. The older taxon Euascomycetes is roughly equivalent.
- The Discomycetes are a subgroup of the Pezizomycotina including cup fungi, morels, etc.
- The subphylum Saccharomycotina comprises the 'true' yeasts which are single-celled and reproduce by budding. These have the older name Hemiascomycetes.
- The disparate subphylum Taphrinomycota includes the remainder of the Ascomycota, which are considered to be more primitive. These can also be known as Archaeoascomycetes.
- Some fungi have been allocated to the Ascomycota on the basis of DNA analysis although they never reproduce sexually and thus have no asci. These are the Mitosporic Ascomycota and form part of the old grouping Deuteromycota (Fungi Imperfecti), a designation which should no longer be needed, since the species can now be related to forms within the main classification system.
Physical make-up
Many of the Ascomycota consist of hyphae, long thin thread-shaped cells often only 5 micrometres thick which form the mycelium, a woolly interlaced mesh. If the hyphae of one mycelium were laid end to end, they could reach a length of several kilometers. At the other extreme are the single-celled yeasts, which can often only be seen with a microscope. A group of species such as for instance the classical Baker's Yeast (Saccharomyces cerevisiae) are however dimorphic, which means that they can appear either in single- or multi-cellular form.
The cell walls of these fungi are almost always formed of Chitin and β-Glucans; individual cells are divided by cross-walls, the septa. These give stability to the hyphae and prevent a great loss of cytoplasm in the event that the cell membrane should be locally damaged. As a result ascomycetes can live in dry environments, contrary to the damp-loving Zygomycota (zygote fungi). Mostly the cell divisions are centrally perforated, so they have a small opening in the middle, through which cytoplasm and also nuclei can move more or less freely throughout the system of hyphae. Most hyphae only have one nucleus per cell, and are therefore described as uninucleate.
The Ascocarp
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In the course of sexual reproduction many, but by no means all, of the ascomycota form macroscopic fruiting bodies which are visible to the naked eye, and which consist of very tightly interwoven hyphae. The fruiting bodies are called ascocarps or ascomas and can sometimes be eaten, for instance in the case of truffles. The ascocarp contains sterile and fertile hyphae, the latter of which give rise to the reproductive cells called spores (in this case ascospores). The reproductive cells are often grouped together in a regular fertile layer, the hymenium, which develops mostly on the inner surface of the ascocarp.
The ascocarp is classified according to its placement (in ways which are not fundamental to the basic taxonomy). It is termed epigeous if it grows above ground, as with the morels, whilst underground ascocarps, such as truffles are hypogeous. The form of the hymenium is divided into the following types.
- Apothecium: here the ascocarp is open above like a cup. The fertile layer is free, so that many spores can be dispersed simultaneously. Morchella (morels), Helvella and Gyromitra have apothecia.
- Cleistothecium: in this case the ascocarp is round with the hymenium enclosed, so the spores do not automatically get released, and fungi with cleistothecia have had to develop new strategies to disseminate their spores. The truffles, for instance, have solved this problem by attracting animals such as wild boars which break open the tasty ascocarps and can spread the spores inside over a wide area. Cleistothecia are found mostly in fungi which have little room available for their ascocarps, for instance those which live under the bark of trees or under the ground like truffles. Also the dermatophyte Arthroderma forms cleistothecia.
- Perithecium: this has the shape of a skittle or a ball. Its distinguishing feature is that on top it has a small pore, the ostiole, through which the spores are released one by one when ripe (in contrast to apothecia where they are released together). Perithecia are found for example on Xylaria (Dead Man’s Fingers, Candle Snuff) and Nectria.
- Pseudothecium: this is similar to a perithecium, but the asci are not regularly organised into a hymenium and they are bitunicate, having a double wall which expands when it takes up water and shoots the enclosed spores out suddenly to disperse them. Example species are Apple scab (Venturia inaequalis) and the horse chestnut disease Guignardia aesculi.
Metabolism
Like most fungi the ascomycota principally digest living or dead biomass. To achieve this, they excrete into their surroundings powerful digestive enzymes which break down organic substances into small molecules, which are then absorbed through the cell wall. Many species live on dead plant material such as fallen leaves, twigs, or indeed large logs. Others attack plants, animals, or other fungi as parasites and derive their metabolic energy, as well as all the nutrients they need, from the cell tissue of their hosts. Especially in this group extreme specialization appears; for instance certain species attack only one particular leg of one particular insect species. The ascomycota also often take up symbiotic relationships - for instance some combine with algae or cyanobacteria, from which they obtain photosynthetic nutrients, to form lichens, and others co-operate with woodland trees as mycorrhizal fungi. Finally there are even carnivorous fungi, which have developed hyphal traps in which they can catch small protists such as amoebae, but also roundworms (Nematoda), rotifers, tardigrades, or even small arthropods such as springtails (Colembola).
In the course of their long evolutionary history the ascomycota have achieved the capability of breaking down almost every organic substance. Unlike nearly all other organisms they are able to digest with their own enzymes plant cellulose and the lignin contained in wood. Also collagen, an important structural protein in animals, and keratin (which hair is made of), serve as food sources. Exotic examples are given by the ascomycete Aureobasidium pullulans, which metabolizes wall paint, and the kerosene fungus Amorphotheca resinae, which (to the misfortune of the airline industry) feeds on aircraft fuel, and in tropical regions sometimes blocks fuel pipes.
Distribution and Living Environment
The Ascomycota are present in all land ecosystems world-wide; they even occur in Antarctica in the form of lichens. On the other hand the distribution of individual species is very variable: some are found on all continents, while for example the white truffle Tuber magnatum, which is much sought after for culinary purposes, only appears regionally in isolated locations of Italy and France.
As already mentioned the septa (cell walls) of the Ascomycota, which divide the cells, enable the colonization of much drier environments than for instance the zygote fungi (Zygomycota). This resistance is extremely marked in the case of certain species which grow on salted fish, which is an effectively very dry situation due to the high osmotic pressure. On the other hand a small group of species has returned to live in water.
Reproduction
The reproduction of the Ascomycota is very varied and can be either asexual or sexual. The reproductive structures which are formed in the latter case, the elongated sac-like asci, are characteristic of the class. On the other hand asexual reproduction plays by far the greater role, and many species have given up the sexual process altogether.
In the Ascomycota two fundamentally different stages in the life cycle can be distinguished: the anamorph state during which the fungus reproduces asexually, and the teleomorph state of sexual reproduction. The combination of both states is sometimes called the holomorph.
Since the anamorph and teleomorph often do not resemble each other superficially at all, until the late 20th century it was often not possible to know that the two stages belonged to the same species. This led to the curious situation, which is still accepted in mycological taxonomy, that two stages in the life cycle of exactly the same fungus have been allocated different species names, so the fungus has two names. For instance the sexual form of the kerosene fungus is known as Amorphotheca resinae while the asexual stage is called Hormoconis resinae. Today however molecular methods lead to the discovery of ever more links between these 'species' and so the former separation of the asexual stages into their own class, the Fungi Imperfecti (Deuteromycota or mitosporic fungi) has now become obsolete.
Asexual reproduction
Image:SEM bread mold (1200x).jpg Asexual reproduction is the dominant form of propagation in the Ascomycota, and is responsible for the rapid expansion of these fungi into areas which were previously not colonized. It occurs through reproductive structures, the conidiospores (or conidia), which are genetically identical to the parent and mostly have just one nucleus. They are also called mitospores due to the way they are generated through the cellular process of mitosis. They are generally formed on the ends of specialized hyphae, the conidiophores. Depending on the species they may be dispersed by wind or water, or also by animals.
Coelomycetes and Hyphomycetes
There is an enormous variety of asexual stages (anamorphs), which can be divided roughly into two groups not having fundamental taxonomic significance, the Coelomycetes and the Hyphomycetes, depending on whether the spores are formed in a closed polyhyphal structure (the conidioma, plural: conidiomata).
- Coelomycetes form their spores in sealed conidiomata which often grow just under the surface of a host organism. One distinguishes:
- acervular conidiomata, or acervuli, which develop in the host and can thus be:
- Mostly they develop a flat layer of relatively short conidiophores which then produce masses of spores. The increasing pressure finally leads to the splitting of the epidermis and cuticle and so allows the conidia to escape.
- pycnidial conidiomata or pycnidia, which unlike the acervuli form in the fungal tissue itself, and which are generally shaped like a bulging vase. The spores are released through a small opening at the apex, the ostiole.
- In the Hyphomycetes the conidiophores (i.e. the hyphae which carry conidia-forming cells on the end) are always free. They are mostly isolated but sometimes also appear as bundles of cells aligned in parallel (described as synnematal) or as cushion-shaped masses (described as sporodochial).
Asexual spores
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In order to further classify the Ascomycota in the asexual stages, it is important to consider the spores, which can be distinguished by colour, form and the way they are separated into cells. The most frequent types are the single-celled spores which are designated amerospores. If the spore is divided into two by a cross-wall (septum), it is a didmyospore. When there are two or more cross-walls the classification depends on the shape. If the septa are transversal, like the rungs of a ladder, it is a phragmospore whilst if they form a net-like structure it is a dictyospore. In staurospores ray-like 'arms' radiate from a central body; in others (helicospores) the entire spore is wound up in a spiral like a spring. Finally very long worm-like spores, of which the ratio length:diameter is more than 15:1, are called scolecospores.
Conidiogenesis and Dehiscence
Two further important characteristics of the anamorphs of the Ascomycota are the conidiogenesis, the fashion in which the spores are formed, and their dehiscence, i.e. how they separate from the parent structures. The former corresponds to Embryology in animals and plants and can be divided into two fundamental forms of development: blastic conidiogenesis, where the spore is already evident before it separates from the conidiogenic hypha which is giving rise to it, and thallic conidiogenesis, where first a cross-wall appears and then the thus created cell develops into a spore.
These two basic types can be further classified as follows.
- blastic-acropetal (repeated budding at the tip of the conidiogenic hypha, so that a chain of spores is formed with the youngest at the tip),
- blastic-synchronous (simultaneous spore formation from a central cell, sometimes with secondary acropetal chains forming from the initial spores),
- blastic-sympodial (repeated sideways spore formation from behind the leading spore, so that the oldest spore is at the main tip),
- blastic-annellidic (each spore separates and leaves a ring-shaped scar which is concentrically inside the scar left by the previous spore),
- blastic-phialidic (the spores arise and are ejected from the open ends of special conidiogenic cells called phialides which remain constant in length; an example is the anamorph of Penicillium),
- basauxic (where a chain of conidia, in successively younger stages of development, is emitted from the mother cell),
- blastic-retrogressive (spores separate off by formation of crosswalls near the tip of the conidiogenic hypha, which thus becomes progressively shorter),
- thallic-arthric (double cell walls split the conidiogenic hypha into cells which develop into short, cylindrical spores called arthroconidia; sometimes every second cell dies off, leaving the arthroconidia free),
- thallic-solitary (a large bulging cell separates from the conidiogenic hypha, forms internal walls, and develops to a phragmospore).
Essentially dehiscence can happen in two different ways. In the schizolytic variant a double dividing wall with a central lamella (layer) forms between the cells; the central layer dissolves to release the spores. In the case of rhexolytic dehiscence on the other hand the cell wall which joins the spores on the outside simply degenerates and sets free the conidia.
Heterocaryosis and Parasexuality
A significant number of Ascomycota species either have no sexual stage or none is known. In spite of this, there are two ways in which they can conserve their genetic diversity: Heterocaryosis and Parasexuality.
The former happens simply through the merging of two hyphae belonging to different individuals, a process known as anastomosis. As a result there are more cell nuclei than normal in the mycelium and they come from genetically different parent organisms.
Parasexuality, on the other hand, refers to a phenomenon where two cell nuclei merge without any sexual process and the chromosome count is doubled. This involves a complex form of the type of cell division called mitosis, where there is crossing over or recombination, i.e. an exchange of genetic material between corresponding pairs of chromosomes. In sexual reproduction, in contrast, crossing over occurs only during meiosis. Finally the chromosome count will be restored to normal by haploidization, whereby the nucleus splits into two parts each having a single set of chromosomes, with each daughter genetically different from the original parents.
Sexual Reproduction
Image:Hypocrea virens.jpg Sexual reproduction in the Ascomycota is marked by a characteristic structure, the ascus, which distinguishes these fungi from all others. An ascus is a tube-shaped vessel, a meiosporangium, which contains the sexual spores produced by meiosis. The latter are called ascospores in contrast to the asexual conidiospores.
Apart from exceptions such as Baker's Yeast (Saccharomyces cerevisiae), almost all fungi of the Ascomycota are haploid, so their nuclei only contain one set of chromosomes, which makes them especially susceptible to mutations. During sexual reproduction there is a diploid phase (with two sets of chromosomes), which as a rule is very short. Then meiosis occurs, generally very soon, so that the haploid state is re-established.
The formation of sexual spores
The sexual part of the life cycle commences when two suitable hyphae meet each other. These come from the same web of hyphae which can also generate asexual spores. The first deciding factor as to whether conjugation - that is, sexual merging - will occur, is whether the hyphae belong to the same organism, or whether they come from different individual fungi. Whilst many species are thoroughly capable of self-propagation, i.e. they are homothallic, others need non-identical partners and so are heterothallic. Besides this, the two hyphae in question must also belong to the same mating type. Mating types are a peculiarity of the fungi and correspond roughly to the sexes in plants and animals; however one species may have more than two mating types.
In the case of compatibility, gametangia form on the hyphae; these are the generative cells for the gametes, in which numerous nuclei gather. A very fine hypha, called the trichogyne, which grows out of one gametangium, now termed the ascogonium, makes a passage to a gametangium of the other indiviual, which is then the antheridium. Nuclei then pass from the antheridium (playing a 'male' role) to the ascogonium (playing a 'female' role).
Unlike the process in animals and plants, after the union of the cytoplasms of the two gametangia (plasmogamy), the merging of the nuclei (karyogamy) does not usually occur immediately. Instead, the nuclei which have migrated in from the antheridium pair up with the nuclei of the ascogonium, but remain separate next to their partners. With this the dikaryophase of the life cycle begins; during this time the pairs of nuclei repeatedly synchronously divide, so that a great number are produced. In all probability the dikaryophase is an evolutionary adaptation which serves to exploit the potential of sexual reproduction to the full in circumstances where it is a rare event for different individuals to meet each other. After the genetic raw material has been increased by repeated division, recombination will take place independently in each pair during meiosis, so that the greatest possible quantity of genetically different spores will arise. In the red algae (Rhodophyta) a similar solution to the corresponding problem evolved independently.
Next millions of new dinucleate hyphae, into each of which two nuclei migrate, emerge from the fertilized ascogonium. They are also called ascogenous or fertile. They are fed by ordinary uni- or mononucleate hyphae (with only one nucleus), which are also called sterile. The tissue of sterile and fertile hyphae now grows in many cases into a macroscopically visible fruiting body, the ascocarp, which may contain millions of fertile hyphae.
In the actual fruiting layer, the hymenium, the asci now appear. At one end of an ascogenous hypha, there develops a U-shaped hook, which points back opposite to the general growth direction. The two nuclei contained in the terminal cell then divide in such a way that the threads of their mitotic spindles run parallel, and thus two pairs of genetically different daughter nuclei arise, with one daughter of each pair near the point of the hook, and the other in the base part of the hypha. Then two parallel cross-walls appear, dividing the hypha into three sections: that at the point of the hook with one nucleus, that at the base of the original hypha with one nucleus, and the middle U-shaped part with two nuclei.
If the positioning in the fruiting layer is right, the karyogamic fusion of the nuclei finally takes place in the U-shaped cell, creating the diploid zygote. It lengthens to form an elongated tube-shaped or cylinder-shaped capsule, the actual ascus. Then meiosis occurs, giving rise to four haploid nuclei. This is almost almost followed by a further mitotic division, so that the ascus ultimately has eight daughter nuclei. These become enclosed, together with some of the cell plasma, each by their own membranes, and generally with a hard cell wall. Thus the dissemination cells (the ascospores) develop, lying initially like peas in a pod inside the ascus. Later, when an appropriate opportunity presents itself, they are liberated.
Not having flagella, ascospores are disseminated in various other ways: some are spread by wind and with others the ripe ascus breaks open on contact with water to set free the spores. Certain species have evolved regular 'spore cannons' which can eject them up to 30 cm. away. When the spores reach a suitable substrate, they germinate, form new hyphae, and so restart their life cycle, which has come full circle.
Ascus classification
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The form of the ascus, the capsule which contains the sexual spores, is important for classification and is divided into four basic types.
- A unitunicate-operculate ascus has a "lid", the Operculum, which breaks open when the spores ripen and in this way sets them free. Unitunicate-operculate asci only occur in those ascocarps which have apothecia, for instance the morels. 'Unitunicate' means 'single-walled'.
- Instead of an operculum, a unitunicate-inoperculate ascus has an elastic ring which functions like a pressure valve. On ripening it briefly extends and so lets the spores shoot out. This type appears both in apothecia and in perithecia; an example is the illustrated Hypomyces chrysospermus.
- A bitunicate ascus is enclosed in a double wall. This consists of a thin brittle outer shell and a thick elastic inner wall. When the spores are ripe the shell splits open so that the inner wall can take up water. As a consequence this begins to extend with its spores until it protrudes above the rest of the ascocarp so that the spores can escape into free air without being obstructed by the bulk of the fruiting body. Bitunicate asci occur only in pseudothecia and are found for instance in Venturia inaequalis (Apple Scab) and Guignardia aesculi (Brown Leaf Mold of Horse Chestnut).
- Prototunicate asci are mostly spherical in shape and they have no active dispersal mechanism at all. The ripe ascus wall simply dissolves so that the spores can escape, or it is broken open by other influences such as animals. Asci of this type can be found both in perithecia and in cleistothecia, for instance with Dutch elm disease (Ophiostoma). This is something of a catch-all term for cases which do not fit into the other three ascus types, and they probably belong to several independent groups which evolved separately from unitunicate asci.
Ecology
The Ascomycota fulfil a central role in most land-based ecosystems. They are important decomposers which break down such organic materials as dead leaves, twigs, fallen trees, etc. and help the detritivores (animals which live off this decomposing material) to obtain their nutrients. By processing substances like cellulose or lignin, which are otherwise difficult to exploit, they take on an important place in the natural nitrogen cycle and the carbon cycle.
Inversely the fruiting bodies of the Ascomycota provide food for a very diverse set of animals from insects and slugs and snails (Gastropoda) to rodents and larger mammals such as deer and wild boars.
Fungi of the Ascomycota are also known for their numerous symbiotic relationships with other organisms.
Lichens
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Probably since early in their evolutionary history the Ascomycota have "domesticated" green algae (Chlorophyta), as well as occasionally other types of algae and cyanobacteria. Together they form the mutualistic associations known as lichens, which can survive in the least hospitable regions of the earth, including the Arctic, the Antarctic, deserts and mountaintops, and can withstand temperature extremes from -40°C to +80°C. While the photoautotrophic algal partner creates metabolic energy through photosynthesis, the fungus offers a stable supportive framework and protects from radiation and drying out. Around 42% of the Ascomycota (numerically about 18,000 species) form lichens, and almost all the fungal partners of lichens belong to the Ascomycota - the proportion of Basidiomycota is probably only two to three percent.
Mycorrhizal fungi and endophytes
Members of the Ascomycota make two particularly important types of relationship with plants: as mycorrhizal fungi and as endophytes. The former make symbiotic associations with the root systems of the plants, which for some trees, especially conifers, can be of vital importance, enabling the uptake of mineral salts from the soil. The fungal partner is in a much better position to absorb minerals due to its finely divided mycelium, whilst the plant provides it with metabolic energy in the form of photosynthetic products. Cases are even known where mycorrhizal fungi can transport nutrients from one plant to another, stabilizing the recipient. It is likely that mycorrhizal associations enabled the conquest of the land by plants - in any case the earliest known fossils of land plants have mycorrhizae.
Endophytes on the other hand live inside plants, especially in the stem and leaves, but generally do not damage their hosts. The exact nature of the relationship between endophytic fungus and host is not yet well understood, but it seems that this form of colonization can bestow a higher resistance against insects, roundworms (nematodes), and bacteria; also it can enable or augment the production of poisonous alkaloids, chemicals which can affect the health of plant-eating mammals.
Symbiotic relationships with animals
A series of Ascomycota species from the genus Xylaria are found in the nests of leafcutter ants and other fungus-growing ants of the tribe Attini and in the fungal gardens of termites (Isoptera). Since they do not generate fruiting bodies until the insects have left the nests, it is suspected that, as confirmed in several cases of Basidiomycota species, they may be cultivated.
On the other hand bark beetles (Scolytidae) are certainly important symbiotic partners. The female beetles transport the spores to new hosts in characteristic tucks in their skin, the mycetangia. There they eat tunnels in the wood, which lead into large chambers in which they lay their eggs. At this time the spores are released and give rise to hyphae which unlike the beetles can digest the wood. The beetle larvae feed on the fungus and after they have metamorphosed into the adult state they again carry spores with them to renew the cycle of infection. A well-known example of this is Dutch elm disease, caused by fungus Ophiostoma ulmi, being carried by the European elm bark beetle Scolytus multistriatus.
Importance for humans
Image:Ophiostoma minus.jpeg Ascomycetes make many contributions to the good of mankind, and also have many ill effects.
Harmful interactions
One of their most harmful roles is as the agent of many plant diseases. For instance:
- Dutch Elm Disease, caused by the closely related species Ophiostoma ulmi and Ophiostoma novo-ulmi, has led to the death of many elms in Europe and North America,
- the originally Asian Cryphonectria parasitica is responsible for attacking Sweet Chestnuts (Castanea sativa"),
- a disease of Maize (Zea mays), which is especially prevalent in North America, is brought about by Cochliobolus heterostrophus, and
- Taphrina deformans causes leaf curl of peach.
Notes
- Template:Note label Template:Harv and reference 4.
- Template:Note label The taxonomic system used here (based on reference 1) is only one among several; another authoratitive one is given by references 2 & 3.
- Template:Note label See reference 5.
References
- All but the first 2 sections are translated from the German article.
- {{Harvard reference
| Surname1=Taylor | First1=John W. | Surname2=Spatafora | First2=Joey | Surname3=Berbee | First3=Mary | Year=1996 | Title=Ascomycota | Periodical=Tree of Life | Date=March 11, 1996 | URL=http://tolweb.org/tree?group=Ascomycota&contgroup=Fungi
}}.
- Ainsworth and Bisby's Dictionary of the Fungi, 9th Edition, see here for more details.
- See the Index Fungorum (Hierarchy Search) for a web search based on the previous reference.
- Outline of Ascomycota - 2001, which is part of MYCONET.
- Palæos Fungi
- C. J. Alexopoulos, M. Blackwell, C. W. Mims: Introductory Mycology, 4th Ed., 1996, ISBN 0-47152-229-5
- B. Kendrick: "The Fifth Kingdom, 3rd Ed., 2001, Kapitel 4, ISBN 1-58510-022-6
- G. J. Krieglsteiner: Verbreitungsatlas der Großpilze Deutschlands (West), Volume 2: Schlauchpilze, Ulmer Verlag, 1993
- F. Breitenbach, J. Kränzlin: Pilze der Schweiz, Volume 1, Ascomycetes, Mykologia Luzern, 1984
- Anamorph-Teleomorph-Datenbank
- Klassifikation aller gültigen Gattungen und höheren Taxa der Schlauchpilze
- Fotografien einiger Fruchtkörper (Ascomata)Template:Link FA
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