Hysteriaceae
Encyclopedia
The Hysteriaceae is a family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 of fungi in the order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 Hysteriales
Hysteriales
The Hysteriales is an order of fungi in the class Dothideomycetes, subclass Pleosporomycetidae. It consists of a single family, Hysteriaceae. Members of Hysteriales produce elongated, often boat shaped sexual structures with slit-like openings . However species with these structures are very diverse...

. They are characterised by having sexual structures called hysterothecia, an elongated structure that opens by a longitudinal slit and releases the sexually produced spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

s. Species of the family have a widespread distribution, especially in temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

 regions, and most are saprobic on wood and bark, although a few are parasitic on plants.

Description

The defining feature of this group, the hysterothecium is a dense, persistent darkly colored structure, with a boat-like shape and a pronounced lengthwise slit. Hysterothecia are capable of opening partially to reveal a lenticular, disk-like hymenium
Hymenium
The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some cells develop into sterile cells called cystidia or...

 or closing tightly in response to relative humidity
Relative humidity
Relative humidity is a term used to describe the amount of water vapor in a mixture of air and water vapor. It is defined as the partial pressure of water vapor in the air-water mixture, given as a percentage of the saturated vapor pressure under those conditions...

. They can be embedded in the substratum, bursting through the surface of the substratum (erumpent), or entirely superficial. They can be solitary or gregarious, ellipsoid to greatly elongated, and are sometimes branched, triradiate or borne on a crust- or net-like growth of mycelium
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...

 (a subiculum). In vertical section, hysterothecia are globose to inversely ovoid (obovoid), with a thick three layered peridium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...

, composed of small pseudoparenchyma
Parenchyma
Parenchyma is a term used to describe a bulk of a substance. It is used in different ways in animals and in plants.The term is New Latin, f. Greek παρέγχυμα - parenkhuma, "visceral flesh", f. παρεγχεῖν - parenkhein, "to pour in" f. para-, "beside" + en-, "in" + khein, "to pour"...

tous cells, the outer layer heavily encrusted with pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

 and often longitudinally striated in age, the middle layer lighter in pigmentation and the inner layer distinctly thin-walled, pallid and compressed.

The hamathecium (hyphae or other tissues between asci) is composed of persistent cellular pseudoparaphyses (hyphae originating above the level of the asci and growing downwards between the developing asci), in a gel matrix, with tips often darkened or branched at maturity to form an epithecium (the external layer of tissue of the fruiting body of lichens and fungi, formed by the union of the tips of the paraphyses over the spore sacs). Bitunicate (double-walled) asci are borne in a basal layer and at maturity are typically club-shaped to cylindrical, bearing eight ascospores, overlapping in two series, ranging from hyaline
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...

 to dark brown, obovoid, clavate, ellipsoid or fusoid. Ascospores are highly diverse in septation
Septate
Septate is a morphological term used in biology in two different instances.*In human biology, it is used to describe a divided uterus.*In mycology, it is used to describe the condition of spores or hyphae in terms of having or lacking a septa to divide...

, and range from didymospores to phragmospores to dityospores, at times surrounded by a gel coating, and often show bipolar asymmetry.
The Hysteriaceae are panglobal in distribution and are primarily lignicolous or corticolous (living on bark), although recently a saxicolous and apparently lichen
Lichen
Lichens are composite organisms consisting of a symbiotic organism composed of a fungus with a photosynthetic partner , usually either a green alga or cyanobacterium...

ized species has been described from Tasmania
Tasmania
Tasmania is an Australian island and state. It is south of the continent, separated by Bass Strait. The state includes the island of Tasmania—the 26th largest island in the world—and the surrounding islands. The state has a population of 507,626 , of whom almost half reside in the greater Hobart...

.

Classification

Current classification of the Hysteriaceae includes the following genera: Hysterium Tode emend. Fr., Hysterographium Corda emend. de Not., Gloniopsis de Not., Gloniella Sacc., Glonium Muhlenb. ex. Fr., Farlowiella Sacc., and Hysterocarina Zogg, to which has been recently added Actidiographium Vassiljeva. The genera Hysteroglonium Rehm ex Lindau, Hysteropatella Rehm and Pseudoscypha Reid & Prioz. are tentatively included in the Hysteriaceae by Eriksson, where, in addition to these, the genera Encephalographa Massal., Graphyllium Clem., and Hemigrapha (Mull. Arg.) R. Sant. ex D. Hawksw. are included by Kirk et al. Both coelomycetous pynidial states (e.g., Hysteropycnis) and dematiaceous hyphomycetous anamorphs (e.g., Coniosporium, Septonema, Sphaeronema and Sporidesmium) have been described for members of the Hysteriaceae.

Historical

The genus Hysterium, the type genus
Type genus
In biological classification, a type genus is a representative genus, as with regard to a biological family. The term and concept is used much more often and much more formally in zoology than it is in botany, and the definition is dependent on the nomenclatural Code that applies:* In zoological...

 of the family Hysteriaceae, is attributed to Tode (1784), who was the first to apply the name to a group of fungi bearing a pronounced longitudinal slit, for which he gave the common name “Venusschwämme”. Recognizing the transitional nature of the ascoma, Tode later (1791) stated: “Medium hoc genus inter Pezizas and Lichenes”. Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...

 (1801) synonimized the epithet pulicaris, first proposed by Lightfoot (1777) for a variety of Lichen scriptus ß pulicaris, into what was to later become the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 of the genus, namely Hysterium pulicare Pers. ex Fr. Early authors provided remarkably accurate figures of hysteriaceous fungi. Bolton (1789) and Greville
Robert Kaye Greville
Robert Kaye Greville was a Scottish mycologist, bryologist, and botanist. He was an accomplished artist and illustrator of natural history. In addition to science he was interested in political causes like abolitionism, capital punishment, keeping Sunday special and the temperance movement...

 (1825) were the first to provide illustrations of asci and ascospores.

Due to the seemingly transitional nature of the hysterothecium, neither fully open nor closed, hysteriaceous fungi have been placed in the discomycetes
Discomycetes
Discomycetes is a former taxonomic class of Ascomycete fungi which contains all of the cup, sponge, brain, and some club-like fungi. It includes typical cup fungi like the scarlet elf cup and the orange peel fungus, and fungi with fruiting bodies of more unusual shape, such as morels, truffles and...

 and pyrenomycetes about equally by various mycologists throughout the 19th Century. In his Systema Mycologicum, Fries
Elias Magnus Fries
-External links:*, Authors of fungal names, Mushroom, the Journal of Wild Mushrooming.*...

 (1823) initially considered hysteriaceous fungi to belong to the pyrenomycetes and placed them in the order Phacidiacei, but later (1835) placed them in his new class discomycetes, stating: “Transitum sistunt ad Discomycetes, sed discum verum non monstrant.” Chevallier (1826) recognized the unique nature of the hysterothecium and was the first to segregate hysteriaceous fungi into a new order, the Hysterineae, which he considered as pyrenomycetes distinct from Fries’ Phacidiei. Corda
August Carl Joseph Corda
August Carl Joseph Corda was a Czech physician and mycologist.-Early life and education:Corda was born in Reichenberg , Bohemia on November 15, 1809. Corda's father was a textile seller...

 (1842), on the other hand, retained the Phacidiei within the Hysteriaceae, and divided the family into a number of subfamilies. De Notaris (1847) considered the Hysteriaceae to belong to the pyrenomycetes and used spore pigmentation to classify hysteriaceous fungi into the Phaeosporii and the Hyalosporii. Saccardo (1873) initially followed Fries, but later (1874) placed hysteriaceous fungi in the pyrenomycetes, and carried de Notaris’ (1847) spore classification scheme further by dividing the Hysteriaceae into nine sections based on pigmentation and the morphology of spore septation. Ellis and Everhart (1892), in their North American Pyrenomycetes, tentatively included the Hysteriaceae, but stated that they had not at first intended to do so due to the transitional nature of the hysterothecium. In Rabenhorst’s Kryptogamen-Flora, Die Pilze, Rehm (1896) compromised and placed the Hysteriales as an order intermediate between the pyrenomycetes and the discomycetes.

Duby (1862) considered hysteriaceous fungi to belong to the pyrenomycetes and proposed two sections, the Hystériées to include Hysterium, Glonium, and Actidium Fr. among others, and the Lophiées to accommodate Ostreichnion Duby, Mytilinidion Duby and Lophium Fr. Although Duby’s (1862) method of classification, based on dehiscent versus nondehiscent asci, was not followed by subsequent workers, he was the first to propose dividing hysteriaceous fungi into what was later to become two distinct families. However, one hundred years would pass before this distinction was fully recognized.

Although von Höhnel (1918) considered the Hysteriaceae to be pyrenomycetes, he proposed a radical revision of the Hysteriales. Many of the fifty plus genera were removed, thereby leaving a greatly reduced core group to form the Hysteriaceae: Hysterium, Hysterographium, Farlowiella, Gloniella, Gloniopsis, Glonium (including the subgenus Psiloglonium von Höhnel for species of Glonium without subiculum), Bulliardella Sacc. (Paoli) [=Actidium], Mytilidion Sacc. [=Mytilinidion], Ostreion Sacc. [=Ostreichnion], Lophium and Dichaena Fr. The Hysteriaceae were seen as closely related to the Lophiostomataceae
Lophiostomataceae
The Lophiostomataceae are a family of fungi in the order Pleosporales. Taxa have a widespread distribution, especially in temperate regions, and saprobic or necrotrophic on herbaceous and woody stems....

and the two were united into a new order, the “Hysterostomaeae” by von Höhnel.

Modern

Modern attempts at classification have placed the Hysteriaceae in the Pseudosphaeriales, the Dothiorales, the Dothideales
Dothideales
The Dothideales are an order of lichenous fungi.-Description:Taxa in this order are characterized by the absence of a hamathecium in a locule, and formation of ovoid to cylindrical fisstunicate asci , usually in bundles or cluster called...

 and in a separate order the Hysteriales, closely related to the Pleosporales
Pleosporales
The Pleosporales is the largest order in the fungal class Dothideomycetes. By some estimates it contains 23 families, 332 genera and more than 4700 species. The majority of species are saprobes on decaying plant material in fresh water, marine, or terrestrial environments, but several species are...

. The Hysteriales were placed in the subclass Loculoascomycetes by Luttrell (1955), corresponding to the Ascoloculares first proposed by Nannfeldt
John Axel Nannfeldt
John-Axel Nannfeldt , born 18 January 1904 in Trelleborg and deceased 4 November 1985 in Uppsala, was a Swedish botanist and mycologist....

 (1932). Unlike the hymenoascomycetes, the loculoascomycete ascoma originates prior to karyogamy
Karyogamy
Karyogamy is the fusion of pronuclei of two cells, as part of syngamy, fertilization, or true bacterial conjugation.It is one of the two major modes of reproduction in fungi...

 in the dikaryon
Dikaryon
Dikaryon is from Greek, di meaning 2 and karyon meaning nut, referring to the cell nucleus.The dikaryon is a nuclear feature which is unique to some fungi, in which after plasmogamy the two compatible nuclei of two cells pair off and cohabit without karyogamy within the cells of the hyphae,...

, with the correlated character state being the functionally two-walled ascus which ruptures in a fissitunicate (like a Jack-in-the-box
Jack-in-the-box
A jack-in-the-box is a children's toy that outwardly consists of a box with a crank. When the crank is turned, it plays a melody, often "Pop Goes the Weasel". At the end of the tune there is a "surprise", the lid pops open and a figure, usually a clown or jester, pops out of the box...

) fashion.

Luttrell (1951) studied ascomal ontogeny and hamathicial development in Glonium stellatum Mühlenb.:Fr. and concluded that the Hysteriaceae possess the pseudoparaphysate Pleospora-type centrum (all the structures enclosed within the ascocarp), in which cellular, septate pseudoparaphyses grow downwards from the cavity roof, initially anchored at both ends, and occupy the locule prior to the formation of asci. Even though G. stellatum clearly possessed the Pleospora-type centrum, it differed in that the locule was an elongated hystereothecium, and the locule appeared to be formed largely by the dissolution of the sterile centrum parenchyma, as in the Dothidea-developmental type. Luttrell (1953) thus concluded that locule
Locule
A locule is a small cavity or compartment within an organ or part of an organism ....

 formation in G. stellatum, presumably representative of the Hysteriaceae as a whole, is intermediate between the Pleospora and Dothidea types. Initially, Luttrell (1953) was unsure whether the Hysteriaceae justified ordinal
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 status, stating that the elongated hysteriaceous locule alone may not appear to be sufficient for the recognition of a separate order and the dothideaceous nature of the centrum at the earliest stages was not observed in his study. Nevertheless, Luttrell (1955) did finally retain the Hysteriales as a separate order and noted that this order may have phylogenetic relationships basal to the Pleosporales.

Zogg (1962) acknowledged the heterogeneity of the classical Hysteriales and, following Duby (1862), divided hysteriaceous fungi into two families, namely the Hysteriaceae s. str. to include Hysterium, Hysterographium, Gloniopsis, Gloniella, Glonium, Farlowiella, and Hysterocarina, and the Lophiaceae Zogg ex von Arx and Müller [equivalent to Mytilinidiaceae Kirschst., as meant by Barr 1990], to include Actidium, Mytilidion [=Mytilinidion], Lophium and Glyphium Nitsch. ex Lehm. Recent molecular evidence, however, has removed the genus Glyphium to the Chaetothyriales
Chaetothyriales
Chaetothyriales is an order of ascomycetous fungi within the class Eurotiomycetes and within the subphylum Pezizomycotina.-References:*C.J. Alexopolous, Charles W. Mims, M. Blackwell, Introductory Mycology, 4th ed. ISBN 0-471-52229-5...

 in the Eurotiomycetes
Eurotiomycetes
The Eurotiomycetes is a class of ascomycetes within the subphylum Pezizomycotina.Some members of the Eurotiomycetes were previously grouped in the class Plectomycetes.-Nomenclature:...

. Zogg (1962) characterized the Hysteriaceae as having ovoid to elongate thick-walled hysterothecia, with a prominent sunken slit, whereas the Mytilinidiaceae, as the Lophiaceae, were characterized by a thin-walled, fragile mussel-shaped (conchate) or hatchet-shaped (dolabrate) ascoma, standing on edge, with a prominent, crested apex. Zogg (1962) postulated that the two families were unrelated, with the Hysteriaceae connected to the Dothioraceae, and the Mytilinidiaceae showing relationships to the Lophiostomataceae.

Although Luttrell held a very wide concept of the Hysteriales (1973), he did not recognize the family Lophiaceae, instead proposing a subfamily within the Hysteriaceae to accommodate mytilinidiaceous forms. Barr (1979) also originally held a wide view of the Hysteriales, but, unlike Luttrell (1973), maintained the two family distinction. Later, Barr (1983) abandoned the Hysteriales and placed the Hysteriaceae within the Pleosporales due to the presence of cellular pseudoparaphyses, asci borne in a basal rather than peripheral layer and ascospores typically showing bipolar asymmetry. Barr noted (1987) that the relationship of some members of the Hysteriaceae was with the Cucurbitariaceae
Cucurbitariaceae
The Cucurbitariaceae are a family of fungi in the order Pleosporales. Taxa are widespread in temperate regions and are necrotrophic or saprobic on woody plants....

 or with the Pleosporaceae. Finally, Kirk et al. (2001) maintained both the Hysteriaceae and the Mytilinidiaceae in the Hysteriales, but Eriksson (2006) removed the Mytilinidiaceae from the Hysteriales and considered it as Dothideomycetes et 'Chaetothyriomycetes incertae sedis
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...

, leaving the Hysteriaceae as the sole family in the Hysteriales.

The Hysteriaceae may show superficial resemblance with the Patellariaceae Corda (Patellariales Hawksw. & Erikss.). But as Barr (1987) points out, the hamathecium in the Patellariaceae is composed of apically free paraphysoids that form a pseudoepithecium, whereas in the Hysteriaceae the hamathecium is composed of downward-growing pseudoparaphyses that, at maturity, become apically dissociated and may become darkened to form an epithecium. Peridial differences, especially the texture and features related to the peridial base, further separate the two families (Barr 1987). The genus Hysteropatella Rehm is transitional with paraphysoids and a well-developed pseudoepithecium, but the peridium, thickened base of the ascoma and cylindric asci are all features of the Hysteriaceae. Kutorga and Hawksworth (1997) in their revision of the Patellariaceae did not include Hysteropatella. Initial studies using the nuclear small subunit (nuSSU) rDNA
Ribosomal DNA
Ribosomal DNA codes for ribosomal RNA. The ribosome is an intracellular macromolecule that produces proteins or polypeptide chains. The ribosome itself consists of a composite of proteins and RNA. As shown in the figure, rDNA consists of a tandem repeat of a unit segment, an operon, composed of...

 have found that Hysterium pulicare and Hysteropatella clavispora (Peck) Seaver formed a clade
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 with high statistical support, distant from other Dothideomycetes, and thus supporting the inclusion of the genus Hysteropatella within the Hysteriaceae.

More recently, Schoch et al. (2006), using a multigene phylogeny of the Dothideomycetes, based on nu SSU, nu LSU, EF1a & RPB2, provided evidence indicating that hysteriaceous fungi occupy a basal position to a monophyletic Pleosporales
Pleosporales
The Pleosporales is the largest order in the fungal class Dothideomycetes. By some estimates it contains 23 families, 332 genera and more than 4700 species. The majority of species are saprobes on decaying plant material in fresh water, marine, or terrestrial environments, but several species are...

, in agreement with Luttrell (1955). However, it was noted that the hysteriaceous fungi sampled did not form a monophyletic group. Farlowiella carmichaeliana (Berk.) Sacc. was basal to the Pleosporales, but very distant from the other members of the Hysteriales sampled, which were inclusive of Hysterium pulicare, Hysteropatella clavispora, and Hysteropatella elliptica Fr., that clustered together with high statistical support, as perhaps the nucleus of an emerging monophyletic Hysteriales. The nearest association of this Hysteriales core group was with members of the Tubeufiaceae M.E. Barr and the Botryosphaeriaceae Theiss. & P. Syd. The sole mytilinidiaceous member analyzed in this study, Lophium mytilinum (Pers.) Fr., also occupied a basal position to the Pleosporales, but was distant to the core group of Hysteriales and was designated as Pleosporomycetidae incertae sedis.

Taken together, classification emphasizing the transitional nature of the hysterothecium, studies in centrum ontogeny
Ontogeny
Ontogeny is the origin and the development of an organism – for example: from the fertilized egg to mature form. It covers in essence, the study of an organism's lifespan...

 and recent molecular evidence, seems to indicate a basal phylogenetic position of the Hysteriales to the Pleosporales and emphasizes the need for further study of the group.

Genera

  • Acrogenospora
    Acrogenospora
    Acrogenospora is a genus of fungi in the family Hysteriaceae.-External links:*...

    M.B. Ellis (1971) has 7 species, all of which are anamorphs of genus Farlowiella. the type species
    Type species
    In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

     is Acrogenospora sphaerocephala (Berk. & Broome) M.B. Ellis (1971).
  • Encephalographa A. Massal. (1854) contains 6 species. Proposed for placement in the Hysteriaceae in 1990.
  • Farlowiella Sacc. (1891), has 2 species. The type is Farlowiella repanda (A. Bloxam ex Duby) Sacc. (1891), which is synonymous with F. carmichaeliana.
  • Gloniella Sacc. (1883) contain over 70 species, and the type is Gloniella sardoa Sacc. & Traverso (1883)
  • Gloniopsis De Not. (1847) contains roughly 45 species. The type is Gloniopsis decipiens De Not. (1847).
  • Glonium Muhl. (1813) contains over 70 species, with the type species being Glonium stellatum Muhl. (1813).
  • Graphyllium
    Graphyllium
    Graphyllium is a genus of fungi in the family Diademaceae.-External links:*...

    Clem. (1901) has 11 species.
Originally placed in the family "Hypodermiaceae". This species genus has linear, membranaceous, narrow ascomata with a thin slit and ascospores that are brown in color, clathrate, and 3-septate
Septate
Septate is a morphological term used in biology in two different instances.*In human biology, it is used to describe a divided uterus.*In mycology, it is used to describe the condition of spores or hyphae in terms of having or lacking a septa to divide...

. There are over 70 species in this genus, and the type is Graphyllium chloës Clem. (1901).
  • Hysterium Pers. (1797), with roughly 400 species described, is the largest genus of the Hysteriaceae. The type is Hysterium pulicare Ellis.
  • Hysterographium Corda (1842), has Hysterographium elongatum Duby as the type. It has over 100 species.
  • Hysteropatella Rehm (1890) has 8 species, and the type is Hysteropatella prostii (Duby) Rehm (1890).
  • Pseudoscypha J. Reid & Piroz (1966) has one species, the type P. abietis.
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