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{{Short description|Family of fungi}}
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{{Cleanup-jargon|date=December 2007}}
{{Use dmy dates|date=October 2023}}
{{Taxobox
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{{Technical|date=December 2007}}
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| name = Hysteriaceae
{{Automatic taxobox
| image =
| image = 2014-03-15 Hysterium pulicare Pers. - Fr 556475.jpg
| image_caption =
| image_caption = ''[[Hysterium pulicare]]''
| regnum = [[Fungus|Fungi]]
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| taxon = Hysteriaceae
| phylum = [[Ascomycota]]
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| authority = [[Chevall.]] (1826)
| classis = [[Ascomycetes]]
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| subdivision_ranks = Genera
| ordo = [[Hysteriales]]
| subdivision =
| familia = Hysteriaceae
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''[[Acrogenospora]]''<br/>
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| familia_authority = Chevall. (1826)
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''[[Actidiographium]]''<br/>
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| subdivision_ranks = [[genus (biology)|Genera]]
| subdivision = ''[[Acrogenospora]]''<br>
''[[Cleistonium]]''<br/>
''[[Actidiographium]]''<br>
''[[Gloniella]]''<br/>
''[[Encephalographa]]''<br>
''[[Gloniopsis]]''<br/>
''[[Farlowiella]]''<br>
''[[Hypodermopsis]]''<br/>
''[[Gloniella]]''<br>
''[[Hysterium]]''<br/>
''[[Gloniopsis]]''<br>
''[[Hysterobrevium]]''<br/>
''[[Glonium]]''<br>
''[[Hysterocarina]]''<br/>
''[[Graphyllium]]''<br>
''[[Hysteroglonium]]''<br/>
''[[Hemigrapha]]''<br>
''[[Hysteropatella]]''<br/>
''[[Hysterium]]''<br>
''[[Oedohysterium]]''<br/>
''[[Hysterocarina]]''<br>
''[[Ostreichnion]]''<br/>
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''[[Hysteroglonium]]''<br>
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''[[Hysterographium]]''<br>
''[[Hysteropatella]]''<br>
''[[Pseudoscypha]]''
''[[Pseudoscypha]]''
}}
}}

The '''Hysteriaceae''' (also known as Dothideomycetes, Ascomycotina, Eumycota) are a [[family (biology)|family]] of [[fungi]] in the [[order (biology)|order]] [[Hysteriales]]. They are characterised by having [[ascoma]] called '''hysterothecia''', an elongated [[spore]]-bearing structure that opens by a longitudinal slit. Species of the family have a widespread distribution, especially in [[temperate]] regions<ref name="isbn0-85199-827-5">{{cite book |author= Cannon PF, Kirk PM.|title=Fungal families of the world |publisher=CABI |location=Wallingford |year=2007 |pages= |isbn=0-85199-827-5 |oclc= |doi= |accessdate=}}</ref>, and most are saprobic on wood and bark, although a few are plant parasites.<ref name="isbn0-471-52229-5">{{cite book |author=Blackwell, Meredith; Alexopoulos, Constantine John; Mims, Charles W. |title=Introductory mycology |publisher=Wiley |location=New York |year=1996 |pages= |isbn=0-471-52229-5 |oclc= |doi= |accessdate=}}</ref>
The '''Hysteriaceae''' are a taxonomic [[family (biology)|family]] of [[fungi]] and the only [[Extant taxon|extant]] family of the [[order (biology)|order]] [[Hysteriales]]. Members of the Hysteriaceae are defined by the possession of a [[ascoma|sexual structure]] called the '''hysterothecium''', an elongated structure that opens by a longitudinal slit and releases sexually produced [[spore]]s. The family is widely distributed, with many species found in [[temperate]] regions,<ref name="isbn0-85199-827-5">{{cite book |vauthors=Cannon PF, Kirk PM |title=Fungal Families of the World |publisher=CAB International |location=Wallingford, UK |year=2007 |isbn=978-0-85199-827-5}}</ref> and most are [[saprobe|saprobic]] on wood and bark, although a few are [[parasitism|parasitic]] on plants.<ref name="isbn0-471-52229-5">{{cite book |author=Blackwell, Meredith |author2=Alexopoulos, Constantine John |author3=Mims, Charles W. |title=Introductory Mycology |publisher=Wiley |location=New York |year=1996 |isbn=0-471-52229-5}}</ref>


==Description==
==Description==
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The defining feature of this group—the hysterothecium—is a dense, persistent darkly colored structure, with a [[wikt:navicular|boat-like]] shape and a pronounced lengthwise slit. Hysterothecia are capable of opening partially to reveal a [[wikt:lenticular|lenticular]] (lens-shaped), disk-like [[hymenium]] or closing tightly in response to [[relative humidity]]. They can be embedded in the [[substrate (biology)|substratum]], bursting through the surface of the substratum ([[erumpent]]), or rest entirely on the surface. They can be solitary or in groups, [[ellipsoid]] to greatly elongated, and are sometimes branched, triradiate or borne on a crust- or net-like growth of [[mycelium]] (a subiculum).<ref name=Zogg1962>Zogg H. (1962). Die Hysteriaceae ''s. str''. und Lophiaceae. ''Beitr Kryptogamenfl Schweiz'' '''11''': 1–190.</ref> In vertical section, hysterothecia are globose to inversely ovoid (obovoid), with a thick three-layered [[peridium]], composed of small [[pseudoparenchyma]]tous cells, the outer layer heavily encrusted with [[biological pigment|pigment]] and often longitudinally [[wikt:striated|striated]] in age, the middle layer lighter in pigmentation and the inner layer distinctly thin-walled, [[wikt:pallid|pallid]] and compressed.<ref name=Barr1987>Barr ME. (1987). Prodromus to class Loculoascomycetes. Hamilton I. Newell, Inc., Amherst, Massachusetts: M.E. Barr Bigelow. 168 p.</ref>


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The hamathecium (hyphae or other tissues between [[ascus|asci]]) is composed of persistent cellular [[pseudoparaphyses]] ([[hypha]]e 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 [[ascospore]]s, overlapping in two series, ranging from [[hyaline]] to dark brown, obovoid, clavate, ellipsoid or [[wikt:fusiod|fusoid]]. Ascospores are highly diverse in [[septate|septation]], and range from didymospores to phragmospores to dityospores, at times surrounded by a gel coating, and often show bipolar asymmetry.<ref name="Zogg1962" /><ref name="Barr1987" />
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Hysteriaceae are defined by a specialized [[ascoma]] termed the hysterothecium. Hysterothecia are dense, persistent [[carbonaceous]] structures, distinctly [[wikt:navicular|navicular]] in outline, and have a pronounced longitudinal slit running the length of the long axis of the fruitbody. Hysterothecia are capable of opening partially to reveal a [[wikt:lenticular|lenticular]], disk-like [[hymenium]] or closing tightly in response to [[relative humidity]]. They can be embedded in the substratum, bursting through the surface of the substratum ([[wikt:erumpent|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]] (a subiculum).<ref name=Zogg1962>Zogg H. (1962). Die Hysteriaceae ''s. str''. und Lophiaceae. ''Beitr Kryptogamenfl Schweiz'' '''11''': 1-190.</ref> In vertical section, hysterothecia are globose to inversely ovoid (obovoid), with a thick three layered [[peridium]], composed of small pseudo[[parenchyma]]tous cells, the outer layer heavily encrusted with [[pigment]] and often longitudinally [[wikt:striated|striated]] in age, the middle layer lighter in pigmentation and the inner layer distinctly thin-walled, [[wikt:pallid|pallid]] and compressed.<ref name=Barr1987>Barr ME. (1987). Prodromus to class Loculoascomycetes. Hamilton I. Newell, Inc., Amherst, Massachusetts: M.E. Barr Bigelow. 168 p.</ref>
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The Hysteriaceae are panglobal in distribution<ref name="Zogg1962" /><ref name=Amano1983>Amano N. (1983). Saprobic loculoascoycetous fungi from Japan 1. Hysteriaceous fungi. ''Trans mycol Soc Japan'' '''24''': 283–297.</ref><ref name=Checa2007>Checa J, Shoemaker RA, Umaña L. (2007). Some new hysteriaceous fungi from [[Costa Rica]]. ''[[Mycologia]]'' '''99''': 285–290.</ref><ref name=Lee2003>Lee S, Crous PW. (2003). Taxonomy and biodiversity of hysteriaceous ascomycetes in fynbos. ''S Afr J Bot'' '''69''': 480–488.</ref><ref name=Lorenzo1998>Lorenzo LE, Messuti MI. (1998). Noteworthy Hysteriaceae from southern South America. ''Mycol Res'' '''102''': 1101–1107.</ref><ref name=Messuti1997>Messuti MI, Lorenzo LE. (1997). A new species of Hysterium from Patagonia, Argentina. ''Mycol Res'' '''101''': 302–304.</ref><ref name=Messuti2003>Messuti MI, Lorenzo LE. (2003). Notes on the genus Hysterographium (Ascomycota, Hysteriaceae) in southern South America. ''Nova Hedwigia'' '''76''': 451–458.</ref><ref name=Teng1933>Teng SC. 1933. Notes on Hysteriales from China. ''Sinensia'' '''4''': 129–144.</ref><ref name=vanderLinde1992>van der Linde EJ. (1992). Notes on the South African Hysteriaceae (Ascomyctes: Mycotina). ''S Afr J Bot'''''58''': 491–499.</ref><ref name=Vasilyeva1999a>Vasilyeva LN. (1999a). Hysteriaceous fungi in the Russian Far East I. Hysterium. ''Mikologiya i Fitopatologiya'' '''33''': 225–227.</ref><ref name=Vasilyeva1999b>Vasilyeva LN. (1999b). Hysteriaceous fungi in the Russian Far East II. The genus Hysterographium. ''Mikologiya i Fitopatologiya'' '''33''': 297–301.</ref><ref name=Vasilyeva2000>Vasilyeva LN. (2000). Hysteriaceous fungi in the Russian Far East III. Glonium and Actidiographium. ''Mikologiya i Fitopatologiya'' '''34''': 3–5.</ref> and are primarily [[wikt: lignicolous|lignicolous]] or corticolous (living on bark), although recently a [[wikt:saxicolous|saxicolous]] and apparently [[lichen]]ized species has been described from [[Tasmania]].<ref name=Kantvilas1997>Kantvilas G, Coppins BJ. (1997). ''Melaspilea circumserpens'' Nyl. rediscovered and referred to ''Glonium'', with discussion on the provenance of some of Robert Brown’s lichen specimens.''Lichenol''. '''29''': 525–531.</ref>


⚫
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]] to dark brown, obovoid, clavate, ellipsoid or [[wikt:fusiod|fusoid]]. Ascospores are highly diverse in [[septate|septation]], and range from didymospores to phragmospores to dityospores, at times surrounded by a gel coating, and often show bipolar asymmetry.<ref name="Zogg1962" /><ref name="Barr1987" />
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The Hysteriaceae are panglobal in distribution<ref name="Zogg1962" /><ref name=Amano1983>Amano N. (1983). Saprobic loculoascoycetous fungi from Japan 1. Hysteriaceous fungi. ''Trans mycol Soc Japan'' '''24''': 283-297.</ref><ref name=Checa2007>Checa J, Shoemaker RA, Umaña L. (2007). Some new hysteriaceous fungi from [[Costa Rica]]. ''[[Mycologia]]'' '''99''': 285-290.</ref><ref name=Lee2003>Lee S, Crous PW. (2003). Taxonomy and biodiversity of hysteriaceous ascomycetes in fynbos. ''S Afr J Bot'' '''69''': 480-488.</ref><ref name=Lorenzo1998>Lorenzo LE, Messuti MI. (1998). Noteworthy Hysteriaceae from southern South America. ''Mycol Res'' '''102''': 1101-1107.</ref><ref name=Messuti1997>Messuti MI, Lorenzo LE. (1997). A new species of Hysterium from Patagonia, Argentina. ''Mycol Res'' '''101''': 302-304.</ref><ref name=Messuti2003>Messuti MI, Lorenzo LE. (2003). Notes on the genus Hysterographium (Ascomycota, Hysteriaceae) in southern South America. ''Nova Hedwigia'' '''76''': 451-458.</ref><ref name=Teng1933>Teng SC. 1933. Notes on Hysteriales from China. ''Sinensia'' '''4''': 129-144.</ref><ref name=vanderLinde1992>van der Linde EJ. (1992). Notes on the South African Hysteriaceae (Ascomyctes: Mycotina). ''S Afr J Bot'''''58''': 491-499.</ref><ref name=Vasilyeva1999a>Vasilyeva LN. (1999a). Hysteriaceous fungi in the Russian Far East I. Hysterium. ''Mikologiya i Fitopatologiya'' '''33''': 225-227.</ref><ref name=Vasilyeva1999b>Vasilyeva LN. (1999b). Hysteriaceous fungi in the Russian Far East II. The genus Hysterographium. ''Mikologiya i Fitopatologiya'' '''33''': 297-301.</ref><ref name=Vasilyeva2000>Vasilyeva LN. (2000). Hysteriaceous fungi in the Russian Far East III. Glonium and Actidiographium. ''Mikologiya i Fitopatologiya'' '''34''': 3-5.</ref> and are primarily [[wikt: lignicolous|lignicolous]] or corticolous (living on bark), although recently a [[wikt:saxicolous|saxicolous]] and apparently [[lichen]]ized species has been described from [[Tasmania]].<ref name=Kantvilas1997>Kantvilas G, Coppins BJ. (1997). ''Melaspilea circumserpens'' Nyl. rediscovered and referred to ''Glonium'', with discussion on the provenance of some of Robert Brown’s lichen specimens.''Lichenol''. '''29''': 525-531.</ref>
==Classification==
==Classification==
Current classification of the Hysteriaceae<ref name=Eriksson2006>Eriksson OE. (2006). Outline of Ascomycota. ''Myconet'' '''12''': 1-88.</ref> 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<ref name="Eriksson2006" />, 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.<ref name=Kirk2001>Kirk PM, Canon PF, David JC, Stalpers JA. (2001). Dictionary of the Fungi. 9th Ed. CAB International.</ref> Both coelomycetous pynidial states (e.g., ''Hysteropycnis'') and dematiaceous hyphomycetous [[anamorph]]s (e.g., ''Coniosporium'', ''Septonema'', ''Sphaeronema'' and ''Sporidesmium'') have been described for members of the ''Hysteriaceae''.<ref name=Lohman1931>Lohman ML. (1931). A study of Glonium parvulum in culture. ''Pap Mich Acad Sci Arts Letters'' '''13''': 141-156.</ref><ref name=Lohman1933>Lohman ML. 1933. Hysteriaceae: Life histories of certain species. ''Pap Mich Acad Sci Arts Letters'' '''17''': 229-288.</ref><ref name=Lohman1934>Lohman ML. (1934). A cultural and taxonomic study of Hysterium hyalinum. ''Pap Mich Acad Sci Arts Letters'' '''19''': 133-140.</ref>
Current classification of the Hysteriaceae<ref name=Eriksson2006>Eriksson OE. (2006). Outline of Ascomycota. ''Myconet'' '''12''': 1–88.</ref> 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 added ''[[Actidiographium]]'' Vassiljeva. The genera ''[[Hysteroglonium]]'' Rehm ex Lindau, ''[[Hysteropatella]]'' Rehm and ''[[Pseudoscypha]]'' Reid & Prioz. are tentatively included in the ''Hysteriaceae'' by Eriksson.<ref name="Eriksson2006" /> Both coelomycetous pynidial states (e.g., ''[[Hysteropycnis]]'') and dematiaceous hyphomycetous [[anamorph]]s (e.g., ''[[Coniosporium]]'', ''[[Septonema]]'', ''Sphaeronema'' and ''[[Sporidesmium]]'') have been described for members of the ''Hysteriaceae''.<ref name=Lohman1931>Lohman ML. (1931). A study of Glonium parvulum in culture. ''Pap Mich Acad Sci Arts Letters'' '''13''': 141–156.</ref><ref name=Lohman1933>Lohman ML. 1933. Hysteriaceae: Life histories of certain species. ''Pap Mich Acad Sci Arts Letters'' '''17''': 229–288.</ref><ref name=Lohman1934>Lohman ML. (1934). A cultural and taxonomic study of Hysterium hyalinum. ''Pap Mich Acad Sci Arts Letters'' '''19''': 133–140.</ref>

===Historical===
===Historical===
The genus ''Hysterium'', the [[type genus]] of the family Hysteriaceae, is attributed to Tode (1784)<ref name=Tode1784>Tode HJ. (1784). Beschreibung des Venusschwämmes (Hysterium). ''Schrift Berlin Ges Naturf Freunde'' '''5''': 53-55.</ref>, 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''”.<ref name=Tode1791>Tode HJ. (1791). ''Fungi Mecklenburgensis II''.</ref> [[Christian Hendrik Persoon|Persoon]] (1801)<ref name=Persoon1801>Persoon CH. (1801). ''Synopsis methodica Fungorum''.</ref> synonimized the epithet ''pulicaris'', first proposed by [[Lightfoot (biologist)|Lightfoot]] (1777)<ref name=Lightfoot1777>Lightfoot J. 1777. ''Flora Scotica''.</ref> for a variety of ''Lichen scriptus'' ß pulicaris, into what was to later become the [[type species]] of the genus, namely ''Hysterium pulicare'' Pers. ex Fr. Early authors provided remarkably accurate figures of hysteriaceous fungi. Bolton (1789)<ref name=Bolton1789>Bolton J. 1789. An history of fungusses growing about Halifax, Vol III.</ref> and [[Robert Kaye Greville|Greville]] (1825)<ref name=Greville1825>Greville RK. 1825. Scottish Cryptogamic Flora, III. Edinburgh, Scotland.</ref> were the first to provide illustrations of asci and ascospores.
The genus ''Hysterium'', the [[type genus]] of the family Hysteriaceae, is attributed to [[Heinrich Julius Tode]] (1784),<ref name=Tode1784>Tode HJ. (1784). Beschreibung des Venusschwämmes (Hysterium). ''Schrift Berlin Ges Naturf Freunde'' '''5''': 53–55.</ref> 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''”.<ref name=Tode1791>Tode HJ. (1791). ''Fungi Mecklenburgensis II''.</ref> [[Christian Hendrik Persoon|Persoon]] (1801)<ref name=Persoon1801>Persoon CH. (1801). ''Synopsis methodica Fungorum''.</ref> synonymized the epithet ''pulicaris'', first proposed by [[Lightfoot (biologist)|Lightfoot]] (1777)<ref name=Lightfoot1777>Lightfoot J. 1777. ''Flora Scotica''.</ref> for a variety of ''Lichen scriptus'' β pulicaris, into what was to later become the [[type species]] of the genus, namely ''Hysterium pulicare'' Pers. ex Fr. Early authors provided remarkably accurate figures of hysteriaceous fungi. [[James Bolton]] (1789)<ref name=Bolton1789>Bolton J. 1789. An history of fungusses growing about Halifax, Vol III.</ref> and [[Robert Kaye Greville]] (1825)<ref name=Greville1825>Greville RK. 1825. Scottish Cryptogamic Flora, III. Edinburgh, Scotland.</ref> 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]] and [[pyrenomycetes]] about equally by various mycologists throughout the 19th Century.<ref name=Bisby1923>Bisby GR. 1923. The literature on the classification of the Hysteriales. ''Trans Brit Mycol Soc'' '''8''': 176-189.</ref> In his ''Systema Mycologicum'', [[Elias Magnus Fries|Fries]] (1823)<ref name=Fries1823>Fries EM. 1823. Systema Mycologicum, sistens fungorum ordines, genera et species hucusque cognitas, II, pars II:276-620.</ref> 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''.”<ref name=Fries1835>Fries EM. 1835. Corpus Florarum provincialium Sueciae, I. Floram Scanicam.</ref> 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''.<ref name=Chevallier1826>Chevallier FF. 1826. Flore général des environs de Paris, Vol I.</ref> [[August Carl Joseph Corda|Corda]] (1842), on the other hand, retained the ''Phacidiei'' within the ''Hysteriaceae'', and divided the family into a number of subfamilies.<ref name=Corda1842>Corda ACJ. 1842. Icones Fungorum Hucusque Cognitorum, Vol V.</ref> 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''.<ref name=DeNotaris1847>De Notaris CG. 1847. Prime linee di una nuova disposizione de Pirenomiceti Isterini. Giorn Bot Ital 2, part I, fasc. '''7-8''': 5-52.</ref> Saccardo (1873) initially followed Fries<ref name=Saccardo1873>Saccardo PA. 1873. Mycologiae Venetae Specimen. ''Atti Soc Venet Trent Sc Nat, Padova'' '''2''': 155-160.</ref>, but later (1874) placed hysteriaceous fungi in the pyrenomycetes<ref name=Saccardo1874>Saccardo PA. 1874. Conspectus generum pyrenomycetum italicorum. ''Atti Soc Venet Trent Sc Nat, Padova'' '''4''': 77-100.</ref>, 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.<ref name=Saccardo1883>Saccardo PA. 1883. Sylloge Fungorum, omnium hucusque cognitorum. '''2''': 776-783.</ref> 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.<ref name=Ellis1892>Ellis JB, Everhart BM. 1892. The North American Pyrenomycetes. Newfield NJ. 793 p.</ref> In Rabenhorst’s ''Kryptogamen-Flora, Die Pilze'', Rehm (1896) compromised and placed the ''Hysteriales'' as an order intermediate between the pyrenomycetes and the discomycetes.<ref name=Rehm1896>Rehm H. 1896. Ascomyceten: Hysteriaceen und Discomyceten, In: ''L. Rabenhorst’s Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz''. 2nd Ed, Eduard Kummer, Leipzig '''3''': 1-56.</ref>
Due to the seemingly transitional nature of the hysterothecium, neither fully open nor closed, hysteriaceous fungi have been placed in the [[discomycetes]] and [[pyrenomycetes]] about equally by various mycologists throughout the 19th Century.<ref name=Bisby1923>Bisby GR. 1923. The literature on the classification of the Hysteriales. ''Trans Brit Mycol Soc'' '''8''': 176–189.</ref> In his ''Systema Mycologicum'', [[Elias Magnus Fries|Fries]] (1823)<ref name=Fries1823>Fries EM. 1823. Systema Mycologicum, sistens fungorum ordines, genera et species hucusque cognitas, II, pars II:276–620.</ref> 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''.”<ref name=Fries1835>Fries EM. 1835. Corpus Florarum provincialium Sueciae, I. Floram Scanicam.</ref> [[François Fulgis 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''.<ref name=Chevallier1826>Chevallier FF. 1826. Flore général des environs de Paris, Vol I.</ref> [[August Carl Joseph Corda|August Corda]] (1842), on the other hand, retained the ''Phacidiei'' within the ''Hysteriaceae'', and divided the family into a number of subfamilies.<ref name=Corda1842>Corda ACJ. 1842. Icones Fungorum Hucusque Cognitorum, Vol V.</ref> [[Giuseppe 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''.<ref name=DeNotaris1847>De Notaris CG. 1847. Prime linee di una nuova disposizione de Pirenomiceti Isterini. Giorn Bot Ital 2, part I, fasc. '''7–8''': 5–52.</ref> [[Pier Andrea Saccardo]] (1873) initially followed Fries,<ref name=Saccardo1873>Saccardo PA. 1873. Mycologiae Venetae Specimen. ''Atti Soc Venet Trent Sc Nat, Padova'' '''2''': 155–160.</ref> but later (1874) placed hysteriaceous fungi in the pyrenomycetes,<ref name=Saccardo1874>Saccardo PA. 1874. Conspectus generum pyrenomycetum italicorum. ''Atti Soc Venet Trent Sc Nat, Padova'' '''4''': 77–100.</ref> 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.<ref name=Saccardo1883>Saccardo PA. 1883. Sylloge Fungorum, omnium hucusque cognitorum. '''2''': 776–783.</ref> [[Job Bicknell Ellis]] and [[Benjamin Matlack 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.<ref name=Ellis1892>Ellis JB, Everhart BM. 1892. The North American Pyrenomycetes. Newfield NJ. 793 p.</ref> In Rabenhorst’s ''Kryptogamen-Flora, Die Pilze'', [[Heinrich Rehm]] (1896) compromised and placed the ''Hysteriales'' as an order intermediate between the pyrenomycetes and the discomycetes.<ref name=Rehm1896>Rehm H. 1896. Ascomyceten: Hysteriaceen und Discomyceten, In: ''L. Rabenhorst’s Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz''. 2nd Ed, Eduard Kummer, Leipzig '''3''': 1–56.</ref>


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.<ref name=Duby1862>Duby JE. 1862. Mémoire sur la tribu des Hystérinées de la famille des Hypoxylées (Pyrénomycètes). ''Mém Soc Phys et Hist Nat Genève'' '''16''': 15-70.</ref> 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.<ref name="Zogg1962" />
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.<ref name=Duby1862>Duby JE. 1862. Mémoire sur la tribu des Hystérinées de la famille des Hypoxylées (Pyrénomycètes). ''Mém Soc Phys et Hist Nat Genève'' '''16''': 15–70.</ref> 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.<ref name="Zogg1962" />

⚫
Although [[Franz Xaver Rudolf von Höhnel|Franz von Höhnel]] (1918) considered the Hysteriaceae to be pyrenomycetes, he proposed a radical revision of the Hysteriales.<ref name=vonHohnel1918>von Höhnel F. 1918. Mycologische Fragmente, 272. Uber die Hysteriaceen. ''Ann Mycol'' '''16''': 145–154.</ref> 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]]'' and the two were united into a new order, the “Hysterostomaeae” by von Höhnel.<ref name="vonHohnel1918" />


⚫
Although von Höhnel (1918) considered the Hysteriaceae to be pyrenomycetes, he proposed a radical revision of the Hysteriales.<ref name=vonHohnel1918>von Höhnel F. 1918. Mycologische Fragmente, 272. Uber die Hysteriaceen. ''Ann Mycol'' '''16''': 145-154.</ref> 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]]'' and the two were united into a new order, the “Hysterostomaeae” by von Höhnel.<ref name="vonHohnel1918" />
===Modern===
===Modern===


Modern attempts at classification have placed the Hysteriaceae in the ''[[Pseudosphaeriales]]''<ref name=Nannfeldt1932>[[John Axel Nannfeldt|Nannfeldt, J. A. 1932. Studien über die Morphologie und Systematik der nicht-lichenisierten, inoperkulaten Discomyceten. ''Nova Acta Regiae Soc Sci Uppsaliensis IV''. '''8'''(2):1-368.</ref><ref name=Gäumann1949>Gäumann EA. 1949. Die Pilze, Grundzüge ihrer Entwicklungsgeschichte und Morphologie. Birkhäuser. Basel, 382 p.</ref>, the ''[[Dothiorales]]''<ref name=Müller1950>Müller E, von Arx JA. 1950. Einige Aspeckte zur Systematik pseudosphärialer Ascomyceten. ''Ber Schwiz Bot Ges'' '''60''': 329-397.</ref><ref name=vonArx1954>von Arx JA, Müller E. 1954. Die Gattungen der amerosporen Pyrenomyceten. ''Beitr Kryptogamenfl Schweiz'' '''11''': 1-434.</ref>, the [[Dothideales]]<ref name=vonArx1975>von Arx JA, Müller E. 1975. A re-evaluation of the bitunicate Ascomycetes with keys to families and genera. ''Stud Mycol Baarn'' '''9''': 1-159.</ref> and in a separate order the ''Hysteriales'', closely related to the [[Pleosporales]].<ref name=Miller1949>Miller JH. 1949. A revision of the classification of the Ascomycetes with special emphasis on the Pyrenomycetes. ''[[Mycologia]]'' '''41''': 99-127.</ref><ref name=Luttrell1955>Luttrell ES. 1955. The ascostromatic ascomycetes. ''[[Mycologia]]'' '''47''': 511-532.</ref> The Hysteriales were placed in the subclass Loculoascomycetes by Luttrell (1955), corresponding to the Ascoloculares first proposed by Nannfeldt (1932). Unlike the hymenoascomycetes, the loculoascomycete ascoma originates prior to [[karyogamy]] in the [[dikaryon]], with the correlated character state being the functionally two-walled ascus which ruptures in a fissitunicate (like a [[Jack-in-the-box]]) fashion.
Modern attempts at classification have placed the Hysteriaceae in the ''[[Pseudosphaeriales]]'',<ref name=Nannfeldt1932>[[John Axel Nannfeldt|Nannfeldt, J. A.]] 1932. Studien über die Morphologie und Systematik der nicht-lichenisierten, inoperkulaten Discomyceten. ''Nova Acta Regiae Soc Sci Uppsaliensis IV''. '''8'''(2):1–368.</ref><ref name="Gäumann1949">Gäumann EA. 1949. Die Pilze, Grundzüge ihrer Entwicklungsgeschichte und Morphologie. Birkhäuser. Basel, 382 p.</ref> the [[Dothiorales]],<ref name="Müller1950">Müller E, von Arx JA. 1950. Einige Aspeckte zur Systematik pseudosphärialer Ascomyceten. ''Ber Schwiz Bot Ges'' '''60''': 329–397.</ref><ref name=vonArx1954>von Arx JA, Müller E. 1954. Die Gattungen der amerosporen Pyrenomyceten. ''Beitr Kryptogamenfl Schweiz'' '''11''': 1–434.</ref> the [[Dothideales]]<ref name=vonArx1975>von Arx JA, Müller E. 1975. A re-evaluation of the bitunicate Ascomycetes with keys to families and genera. ''Stud Mycol Baarn'' '''9''': 1–159.</ref> and in a separate order the ''Hysteriales'', closely related to the [[Pleosporales]].<ref name=Miller1949>Miller JH. 1949. A revision of the classification of the Ascomycetes with special emphasis on the Pyrenomycetes. ''[[Mycologia]]'' '''41''': 99–127.</ref><ref name=Luttrell1955 /> The Hysteriales were placed in the subclass Loculoascomycetes by Luttrell (1955), corresponding to the Ascoloculares first proposed by [[John Axel Nannfeldt|Nannfeldt]] (1932). Unlike the hymenoascomycetes, the loculoascomycete ascoma originates prior to [[karyogamy]] in the [[dikaryon]], with the correlated character state being the functionally two-walled ascus which ruptures in a [[fissitunicate]] (like a [[Jack-in-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.<ref name=Luttrell1951>Luttrell ES. 1951. Taxonomy of the Pyrenomycetes. ''Univ Missouri Stud Sci'' '''24''': 1-120.</ref> 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]] formation in ''G. stellatum'', presumably representative of the Hysteriaceae as a whole, is intermediate between the ''Pleospora'' and ''Dothidea'' types<ref name=Luttrell1953>Luttrell ES. 1953. Development of the ascocarp in Glonium stellatum. ''Amer J Bot'' '''40''': 626-633.</ref>. Initially, Luttrell (1953) was unsure whether the Hysteriaceae justified [[Order (biology)|ordinal]] 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.<ref name="Luttrell1955" />
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.<ref name=Luttrell1951>Luttrell ES. 1951. Taxonomy of the Pyrenomycetes. ''Univ Missouri Stud Sci'' '''24''': 1–120.</ref> 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]] formation in ''G. stellatum'', presumably representative of the Hysteriaceae as a whole, is intermediate between the ''Pleospora'' and ''Dothidea'' types.<ref name=Luttrell1953>Luttrell ES. 1953. Development of the ascocarp in Glonium stellatum. ''Amer J Bot'' '''40''': 626–633.</ref> Initially, Luttrell (1953) was unsure whether the Hysteriaceae justified [[Order (biology)|ordinal]] 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 (phylogenetics)|basal]] to the Pleosporales.<ref name="Luttrell1955" />


Zogg (1962) acknowledged the heterogeneity of the classical Hysteriales<ref name="Zogg1962" /> and, following Duby (1862)<ref name="Duby1862" />, 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<ref name="Zogg1962" /><ref name="vonArx1975" /> [equivalent to Mytilinidiaceae Kirschst., as meant by Barr 1990<ref name=Barr1990>Barr ME. 1990. Melanomatales (Loculoascomycetes). North American Flora, Series II, Part 13: 1-129.</ref>], to include ''Actidium'', ''Mytilidion'' [=''Mytilinidion''], ''Lophium'' and ''Glyphium'' Nitsch. ex Lehm. Recent molecular evidence, however, has removed the genus ''Glyphium'' to the [[Chaetothyriales]] in the [[Eurotiomycetes]].<ref name=Lindemuth2001>Lindemuth R, Wirtz N, Lumbsch HT. 2001. Phylogenetic analysis of nuclear and mitochondrial rDNA sequences supports the view that loculoascomycetes (Ascomyctoa) are not monophyletic. ''Mycol Res'''''105''': 1176-1181.</ref><ref name=Kodsueb2006>Kodsueb R, Dhanasekaran V, Aptroot A, Lumyong S, McKenzie EHC, Hyde KD, Jeewon R. 2006. The family Pleosporaceae: intergeneric relationships and phylogenetic perspectives based on sequence analyses of partial 28S rDNA. ''[[Mycologia]]'' '''98''': 571-583.</ref><ref name=Geiser2006>Geiser D, Gueidan C, Miadlikowska J, Lutzoni F, Kauff F, Hofstetter V, Fraker E, Schoch CL, Tibell L. 2006. Eurotiomycetes: Eurotiomycetidae and Chaetthyriomycetidae.''[[Mycologia]]'' '''98''': 1053-1064.</ref> 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.
Hans Zogg (1962) acknowledged the heterogeneity of the classical Hysteriales<ref name="Zogg1962" /> and, following Duby (1862),<ref name="Duby1862" /> 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<ref name="Zogg1962" /><ref name="vonArx1975" /> [equivalent to Mytilinidiaceae Kirschst., as meant by [[Margaret Elizabeth Barr-Bigelow|Barr]] 1990<ref name=Barr1990>Barr ME. 1990. Melanomatales (Loculoascomycetes). North American Flora, Series II, Part 13: 1–129.</ref>], to include ''Actidium'', ''Mytilidion'' [=''Mytilinidion''], ''Lophium'' and ''Glyphium'' Nitsch. ex Lehm. Recent molecular evidence, however, has removed the genus ''Glyphium'' to the [[Chaetothyriales]] in the [[Eurotiomycetes]].<ref name=Lindemuth2001>Lindemuth R, Wirtz N, Lumbsch HT. 2001. Phylogenetic analysis of nuclear and mitochondrial rDNA sequences supports the view that loculoascomycetes (Ascomyctoa) are not monophyletic. ''Mycol Res'''''105''': 1176–1181.</ref><ref name=Kodsueb2006>Kodsueb R, Dhanasekaran V, Aptroot A, Lumyong S, McKenzie EHC, Hyde KD, Jeewon R. 2006. The family Pleosporaceae: intergeneric relationships and phylogenetic perspectives based on sequence analyses of partial 28S rDNA. ''[[Mycologia]]'' '''98''': 571–583.</ref><ref name=Geiser2006>Geiser D, Gueidan C, Miadlikowska J, Lutzoni F, Kauff F, Hofstetter V, Fraker E, Schoch CL, Tibell L. 2006. Eurotiomycetes: Eurotiomycetidae and Chaetthyriomycetidae.''[[Mycologia]]'' '''98''': 1053–1064.</ref> 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.<ref name=Luttrell1973>Luttrell ES. 1973. Loculoascomycetes. In Ainsworth GC, Sparrow FK, Sussman AS, eds. ''The Fungi: an advanced treatise''. IVA. London, Academic Press. p. 135-219.</ref> Barr (1979) also originally held a wide view of the Hysteriales<ref name=Barr1979>Barr ME. 1979. A classification of loculoascomycetes. ''[[Mycologia]]'' '''71''': 935-957.</ref>, 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.<ref name=Barr1983>Barr ME. 1983. The ascomycete connection. ''[[Mycologia]]'' '''75''': 1-13.</ref> Barr noted (1987) that the relationship of some members of the Hysteriaceae was with the [[Cucurbitariaceae]] or with the Pleosporaceae.<ref name="Barr1987" /> Finally, Kirk et al. (2001) maintained both the ''Hysteriaceae'' and the ''Mytilinidiaceae'' in the ''Hysteriales''<ref name="Kirk2001" />, but Eriksson (2006) removed the Mytilinidiaceae from the Hysteriales and considered it as Dothideomycetes et 'Chaetothyriomycetes ''[[incertae sedis]]'', leaving the Hysteriaceae as the sole family in the Hysteriales.<ref name="Eriksson2006" />
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.<ref name=Luttrell1973>Luttrell ES. 1973. Loculoascomycetes. In Ainsworth GC, Sparrow FK, Sussman AS, eds. ''The Fungi: an advanced treatise''. IVA. London, Academic Press. p. 135–219.</ref> Barr (1979) also originally held a wide view of the Hysteriales,<ref name=Barr1979>Barr ME. 1979. A classification of loculoascomycetes. ''[[Mycologia]]'' '''71''': 935–957.</ref> 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.<ref name=Barr1983>Barr ME. 1983. The ascomycete connection. ''[[Mycologia]]'' '''75''': 1–13.</ref> Barr noted (1987) that the relationship of some members of the Hysteriaceae was with the [[Cucurbitariaceae]] or with the Pleosporaceae.<ref name="Barr1987" /> Finally, Kirk et al. (2001) maintained both the ''Hysteriaceae'' and the ''Mytilinidiaceae'' in the ''Hysteriales'',<ref name="Kirk2001">{{cite book |vauthors=Kirk PM, Cannon PF, David JC, Stalpers JA |title=Ainsworth & Bisby's Dictionary of the Fungi |edition=9th |publisher=CABI Bioscience |location=Oxon, UK |year=2001 |isbn=978-0-85199-377-5}}</ref> but Eriksson (2006) removed the Mytilinidiaceae from the Hysteriales and considered it as Dothideomycetes et 'Chaetothyriomycetes ''[[incertae sedis]]'', leaving the Hysteriaceae as the sole family in the Hysteriales.<ref name="Eriksson2006" />


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 appically dissociated and may become darkened to form an epithecium.<ref name="Barr1987" /> Peridial differences, especially the texture and features related to the peridial base, further separate the two families (Barr 1987).<ref name="Barr1987" /> The genus ''Hysteropatella'' Rehm is transitional with paraphysoids and a well-developed pseudoepithecium, but the peridium, thickend base of the ascoma and cylindric asci are all features of the Hysteriaceae.<ref name="Barr1987" /> Kutorga and Hawksworth (1997) in their revision of the ''Patellariaceae'' did not include Hysteropatella.<ref name=Kutorga1997>Kutorga E, Hawksworth DL. 1997. A reassessment of the genera referred to the family Patellariaceae (Ascomycota). ''Syst Asco'' '''15''': 1-110.</ref> Initial studies using the nuclear small subunit (nuSSU) [[Ribosomal DNA|rDNA]]<ref name=Winka2000>Winka K, Erikson OE. 2000 Adding to the bitunicate puzzle: studies on the systematic positions of five aberrant ascomycete taxa. In: K. Winka. Phylogenetic relationships within the Ascomycota based on 18S rDNA sequences '''5''': 1-13. Umeå (Sweden): PhD Thesis, Umeå University.</ref><ref name=Liew2000>Liew ECY, Aptroot A, Hyde KD. 2000. Phylogenetic significance of the pseudoparaphyses in Loculoascomycete taxonomy. ''Mol Phylogenet Evol'' '''16''': 392-402.</ref> have found that ''Hysterium pulicare'' and ''Hysteropatella clavispora'' (Peck) Seaver formed a [[cladistics|clade]] with high statistical support, distant from other Dothideomycetes, and thus supporting the inclusion of the genus Hysteropatella within the Hysteriaceae.<ref name="Eriksson2006" />
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.<ref name="Barr1987" /> Peridial differences, especially the texture and features related to the peridial base, further separate the two families (Barr 1987).<ref name="Barr1987" /> 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.<ref name="Barr1987" /> Kutorga and Hawksworth (1997) in their revision of the ''Patellariaceae'' did not include Hysteropatella.<ref name=Kutorga1997>Kutorga E, Hawksworth DL. 1997. A reassessment of the genera referred to the family Patellariaceae (Ascomycota). ''Syst Asco'' '''15''': 1–110.</ref> Initial studies using the nuclear small subunit (nuSSU) [[Ribosomal DNA|rDNA]]<ref name=Winka2000>Winka K, Erikson OE. 2000 Adding to the bitunicate puzzle: studies on the systematic positions of five aberrant ascomycete taxa. In: K. Winka. Phylogenetic relationships within the Ascomycota based on 18S rDNA sequences '''5''': 1–13. Umeå (Sweden): PhD Thesis, Umeå University.</ref><ref name=Liew2000>Liew ECY, Aptroot A, Hyde KD. 2000. Phylogenetic significance of the pseudoparaphyses in Loculoascomycete taxonomy. ''Mol Phylogenet Evol'' '''16''': 392–402.</ref> have found that ''Hysterium pulicare'' and ''Hysteropatella clavispora'' (Peck) Seaver formed a [[cladistics|clade]] with high statistical support, distant from other Dothideomycetes, and thus supporting the inclusion of the genus Hysteropatella within the Hysteriaceae.<ref name="Eriksson2006" />


More recently, Schoch et al. (2006)<ref name=Schoch2006>Schoch CL, Shoemaker RA, Seifert KA, Hambleton S. Spatafora JW, Crous PW. 2006. A multigene phylogeny of the Dothideomycetes using four nuclear loci. ''[[Mycologia]]'' '''98''': 1041-1052.</ref>, 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]], in agreement with Luttrell (1955).<ref name=Luttrell1955>Luttrell ES. (1955). The ascostromatic ascomycetes. ''[[Mycologia]]'' '''47''': 511-532.</ref> However, it was noted that the hysteriaceous fungi sampled did not form a monophyletic group.<ref name="Schoch2006" /> ''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.<ref name="Schoch2006" /> 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''.<ref name="Schoch2006" />
More recently, Schoch et al. (2006),<ref name=Schoch2006>Schoch CL, Shoemaker RA, Seifert KA, Hambleton S. Spatafora JW, Crous PW. 2006. A multigene phylogeny of the Dothideomycetes using four nuclear loci. ''[[Mycologia]]'' '''98''': 1041–1052.</ref> 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]], in agreement with Luttrell (1955).<ref name=Luttrell1955>Luttrell ES. (1955). The ascostromatic ascomycetes. ''[[Mycologia]]'' '''47''': 511–532.</ref> However, it was noted that the hysteriaceous fungi sampled did not form a monophyletic group.<ref name="Schoch2006" /> ''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.<ref name="Schoch2006" /> 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''.<ref name="Schoch2006" />


Taken together, classification emphasizing the transitional nature of the hysterothecium, studies in centrum [[ontogeny]] 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.
Taken together, classification emphasizing the transitional nature of the hysterothecium, studies in centrum [[ontogeny]] 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==
==Genera==
*''[[Acrogenospora]]'' M.B. Ellis (1971)<ref name="isbn0-85198-618-8">{{cite book |author=Ellis MB. |title=Dematiaceous Hyphomycetes (Cabi Publishing) |publisher=CABI |location=Wallingford |year=1971 |pages= |isbn=0-85198-618-8 |oclc= |doi= |accessdate=}}</ref> has 7 species, all of which are [[anamorph]]s of genus [[Farlowiella]].<ref name="urlIndex Fungorum - ''Acrogenospora'' species list">{{cite web |url=http://www.indexfungorum.org/Names/names.asp?strGenus=Acrogenospora |title=Index Fungorum - ''Acrogenospora'' species list"|format= |work= |accessdate=2008-12-04}}</ref> the [[type species]] is ''Acrogenospora sphaerocephala'' (Berk. & Broome) M.B. Ellis (1971).
*''[[Acrogenospora]]'' M.B. Ellis (1971)<ref name="isbn0-85198-618-8">{{cite book |author=Ellis MB. |title=Dematiaceous Hyphomycetes (Cabi Publishing) |publisher=CABI |location=Wallingford |year=1971 |isbn=0-85198-618-8 }}</ref> has 7 species, all of which are [[anamorph]]s of genus [[Farlowiella]].<ref name="urlIndex Fungorum - ''Acrogenospora'' species list">{{cite web |url=http://www.indexfungorum.org/Names/names.asp?strGenus=Acrogenospora |title=Index Fungorum – ''Acrogenospora'' species list|access-date=4 December 2008}}</ref> the [[type species]] is ''Acrogenospora sphaerocephala'' (Berk. & Broome) M.B. Ellis (1971).
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*''[[Gloniella]]'' Sacc. (1883) contain over 70 species, and the type is ''Gloniella sardoa'' Sacc. & Traverso (1883)<ref name="urlGenus Record Details:''Gloniella''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2089 |title=Genus Record Details:Gloniella|access-date=4 December 2008}}</ref>
*''[[Encephalographa]]'' A. Massal. (1854) contains 6 species. Proposed for placement in the Hysteriaceae in 1990.<ref>Renobales G, Aguirre B. (1990). The nomenclature and systematic position of the genus ''Encephalographa''. ''Systema Ascomycetum'' '''8'''(2): 87-92.</ref>
*''[[Farlowiella]]'' Sacc. (1891), has 2 species. The type is ''Farlowiella repanda'' (A. Bloxam ex Duby) Sacc. (1891), which is synonymous with ''F. carmichaeliana''.<ref name="urlGenus Record Details:Farlowiella">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=1975 |title=Genus Record Details:Farlowiella|format= |work= |accessdate=2008-12-04}}</ref>
*''[[Gloniopsis]]'' De Not. (1847) contains roughly 45 species. The type is ''Gloniopsis decipiens'' De Not. (1847).<ref name="urlGenus Record Details:''Gloniopsis''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2090 |title=Genus Record Details:Gloniopsis |access-date=4 December 2008}}</ref>
*''[[Gloniella]]'' Sacc. (1883) contain over 70 species, and the type is ''Gloniella sardoa'' Sacc. & Traverso (1883)<ref name="urlGenus Record Details:''Gloniella''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2089 |title=Genus Record Details:Gloniella|format= |work= |accessdate=2008-12-04}}</ref>
*''[[Hysterium]]'' Pers. (1797), with roughly 400 species described, is the largest genus of the Hysteriaceae.<ref name="urlIndex Fungorum - ''Hysterium'' species list">{{cite web |url=http://www.indexfungorum.org/Names/names.asp?strGenus=Hysterium |title=Index Fungorum – ''Hysterium'' species list |access-date=4 December 2008}}</ref> The type is ''Hysterium pulicare'' Ellis.<ref name="urlGenus Record Details:''Hysterium''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2464 |title=Genus Record Details:''Hysterium''|access-date=4 December 2008}}</ref>
*''[[Gloniopsis]]'' De Not. (1847) contains roughly 45 species. The type is ''Gloniopsis decipiens'' De Not. (1847).<ref name="urlGenus Record Details:''Gloniopsis''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2090 |title=Genus Record Details:Gloniopsis |format= |work= |accessdate=2008-12-04}}</ref>
*''[[Hysteropatella]]'' Rehm (1890) has 8 species,<ref name="urlIndex Fungorum - Search Page:''Hysteropatella''">{{cite web |url=http://www.indexfungorum.org/Names/names.asp?strGenus=Hysteropatella |title=Index Fungorum – Search Page:''Hysteropatella''|access-date=4 December 2008}}</ref> and the type is ''Hysteropatella prostii'' (Duby) Rehm (1890).<ref name="urlGenus Record Details:''Hysteropatella''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2469 |title=Genus Record Details:''Hysteropatella'' |access-date=4 December 2008}}</ref>
*''[[Glonium]]'' Muhl. (1813) contains over 70 species, with the type species being ''Glonium stellatum'' Muhl. (1813).<ref name="urlGenus Record Details:''Glonium''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2091 |title=Genus Record Details:''Glonium''|format= |work= |accessdate=2008-12-04}}</ref>
*''[[Pseudoscypha]]'' J. Reid & Piroz (1966) has one species, the type ''P. abietis''.<ref name="urlGenus Record Details:''Pseudoscypha''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=4486 |title=Genus Record Details:''Pseudoscypha'' |access-date=4 December 2008}}</ref>

*''[[Graphyllium]]'' Clem. (1901) has 11 species.
== References ==
: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]].<ref>Shoemaker RA, Babcock CE. (1992). Aplanodictyosporous Pleosporales: ''Clathrospora'', ''Comoclathris'', ''Graphyllium'', ''Macrospora'', and ''Platysporoides''. ''Canadian Journal of Botany'' '''70'''(8): 1617-1658.</ref> There are over 70 species in this genus, and the type is ''Graphyllium chloës'' Clem. (1901).<ref name="urlGenus Record Details:''Graphyllium''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2133 |title=Genus Record Details:''Graphyllium''|format= |work= |accessdate=2008-12-04}}</ref>
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{{Reflist|colwidth=30em}}
*''[[Hysterium]]'' Pers. (1797), with roughly 400 species described, is the largest genus of the Hysteriaceae.<ref name="urlIndex Fungorum - ''Hysterium'' species list">{{cite web |url=http://www.indexfungorum.org/Names/names.asp?strGenus=Hysterium |title=Index Fungorum - ''Hysterium'' species list |format= |work= |accessdate=2008-12-04}}</ref> The type is ''Hysterium pulicare'' Ellis.<ref name="urlGenus Record Details:''Hysterium''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2464 |title=Genus Record Details:''Hysterium''|format= |work= |accessdate=2008-12-04}}</ref>
*''[[Hysterographium]]'' Corda (1842), has ''Hysterographium elongatum'' Duby as the type.<ref name="urlGenus Record Details:''Hysterographium''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2468 |title=Genus Record Details:''Hysterographium''|format= |work= |accessdate=2008-12-04}}</ref> It has over 100 species.<ref name="urlIndex Fungorum - Search Page:''Hysterographium''">{{cite web |url=http://www.indexfungorum.org/Names/names.asp?strGenus=Hysterographium |title=Index Fungorum - Search Page:''Hysterographium''|format= |work= |accessdate=2008-12-04}}</ref>
*''[[Hysteropatella]]'' Rehm (1890) has 8 species<ref name="urlIndex Fungorum - Search Page:''Hysteropatella''">{{cite web |url=http://www.indexfungorum.org/Names/names.asp?strGenus=Hysteropatella |title=Index Fungorum - Search Page:''Hysteropatella''|format= |work= |accessdate=2008-12-04}}</ref>, and the type is ''Hysteropatella prostii'' (Duby) Rehm (1890).<ref name="urlGenus Record Details:''Hysteropatella''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=2469 |title=Genus Record Details:''Hysteropatella'' |format= |work= |accessdate=2008-12-04}}</ref>
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*''[[Pseudoscypha]]'' J. Reid & Piroz (1966) has one species, the type ''P. abietis''.<ref name="urlGenus Record Details:''Pseudoscypha''">{{cite web |url=http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=4486 |title=Genus Record Details:''Pseudoscypha'' |format= |work= |accessdate=2008-12-04}}</ref>


==References==
==Other sources==
* Boehm EWA, Schoch CL, Spatafora JW (2009). On the evolution of the Hysteriaceae and Mytilinidiaceae (Pleosporomycetidae, Dothideomycetes, Ascomycota) using four nuclear genes. ''Mycol Res'' '''113''': 461–479.
* Boehm EWA, Mugambi GK, Miller AN, Huhndorf SM, Marincowitz S, Spatafora JW, Schoch CL. (2009). A molecular phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae, and Gloniaceae (Pleosporomycetidae, Dothideomycetes) with keys to world species. ''Studies in Mycology'' '''64''': 49 – 83.


{{Taxonbar|from=Q5962348}}
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{{Reflist}}
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[[Category:Dothideomycetes]]
[[Category:Hysteriales]]
[[Category:Ascomycota families]]
[[Category:Taxa described in 1826]]
[[Category:Taxa named by François Fulgis Chevallier]]

Latest revision as of 00:50, 15 May 2024

Hysteriaceae
Hysterium pulicare
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Dothideomycetes
Order: Hysteriales
Family: Hysteriaceae
Chevall. (1826)
Genera

Acrogenospora
Actidiographium
Cleistonium
Gloniella
Gloniopsis
Hypodermopsis
Hysterium
Hysterobrevium
Hysterocarina
Hysteroglonium
Hysteropatella
Oedohysterium
Ostreichnion
Pseudoscypha

The Hysteriaceae are a taxonomic family of fungi and the only extant family of the order Hysteriales. Members of the Hysteriaceae are defined by the possession of a sexual structure called the hysterothecium, an elongated structure that opens by a longitudinal slit and releases sexually produced spores. The family is widely distributed, with many species found in temperate regions,[1] and most are saprobic on wood and bark, although a few are parasitic on plants.[2]

Description

[edit]

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 (lens-shaped), disk-like hymenium or closing tightly in response to relative humidity. They can be embedded in the substratum, bursting through the surface of the substratum (erumpent), or rest entirely on the surface. They can be solitary or in groups, ellipsoid to greatly elongated, and are sometimes branched, triradiate or borne on a crust- or net-like growth of mycelium (a subiculum).[3] In vertical section, hysterothecia are globose to inversely ovoid (obovoid), with a thick three-layered peridium, composed of small pseudoparenchymatous cells, the outer layer heavily encrusted with pigment and often longitudinally striated in age, the middle layer lighter in pigmentation and the inner layer distinctly thin-walled, pallid and compressed.[4]

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 to dark brown, obovoid, clavate, ellipsoid or fusoid. Ascospores are highly diverse in septation, and range from didymospores to phragmospores to dityospores, at times surrounded by a gel coating, and often show bipolar asymmetry.[3][4] The Hysteriaceae are panglobal in distribution[3][5][6][7][8][9][10][11][12][13][14][15] and are primarily lignicolous or corticolous (living on bark), although recently a saxicolous and apparently lichenized species has been described from Tasmania.[16]

Classification

[edit]

Current classification of the Hysteriaceae[17] 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 added Actidiographium Vassiljeva. The genera Hysteroglonium Rehm ex Lindau, Hysteropatella Rehm and Pseudoscypha Reid & Prioz. are tentatively included in the Hysteriaceae by Eriksson.[17] 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.[18][19][20]

Historical

[edit]

The genus Hysterium, the type genus of the family Hysteriaceae, is attributed to Heinrich Julius Tode (1784),[21] 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”.[22] Persoon (1801)[23] synonymized the epithet pulicaris, first proposed by Lightfoot (1777)[24] for a variety of Lichen scriptus β pulicaris, into what was to later become the type species of the genus, namely Hysterium pulicare Pers. ex Fr. Early authors provided remarkably accurate figures of hysteriaceous fungi. James Bolton (1789)[25] and Robert Kaye Greville (1825)[26] 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 and pyrenomycetes about equally by various mycologists throughout the 19th Century.[27] In his Systema Mycologicum, Fries (1823)[28] 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.”[29] François Fulgis 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.[30] August Corda (1842), on the other hand, retained the Phacidiei within the Hysteriaceae, and divided the family into a number of subfamilies.[31] Giuseppe 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.[32] Pier Andrea Saccardo (1873) initially followed Fries,[33] but later (1874) placed hysteriaceous fungi in the pyrenomycetes,[34] 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.[35] Job Bicknell Ellis and Benjamin Matlack 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.[36] In Rabenhorst’s Kryptogamen-Flora, Die Pilze, Heinrich Rehm (1896) compromised and placed the Hysteriales as an order intermediate between the pyrenomycetes and the discomycetes.[37]

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.[38] 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.[3]

Although Franz von Höhnel (1918) considered the Hysteriaceae to be pyrenomycetes, he proposed a radical revision of the Hysteriales.[39] 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 and the two were united into a new order, the “Hysterostomaeae” by von Höhnel.[39]

Modern

[edit]

Modern attempts at classification have placed the Hysteriaceae in the Pseudosphaeriales,[40][41] the Dothiorales,[42][43] the Dothideales[44] and in a separate order the Hysteriales, closely related to the Pleosporales.[45][46] The Hysteriales were placed in the subclass Loculoascomycetes by Luttrell (1955), corresponding to the Ascoloculares first proposed by Nannfeldt (1932). Unlike the hymenoascomycetes, the loculoascomycete ascoma originates prior to karyogamy in the dikaryon, with the correlated character state being the functionally two-walled ascus which ruptures in a fissitunicate (like a Jack-in-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.[47] 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 formation in G. stellatum, presumably representative of the Hysteriaceae as a whole, is intermediate between the Pleospora and Dothidea types.[48] Initially, Luttrell (1953) was unsure whether the Hysteriaceae justified ordinal 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.[46]

Hans Zogg (1962) acknowledged the heterogeneity of the classical Hysteriales[3] and, following Duby (1862),[38] 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[3][44] [equivalent to Mytilinidiaceae Kirschst., as meant by Barr 1990[49]], to include Actidium, Mytilidion [=Mytilinidion], Lophium and Glyphium Nitsch. ex Lehm. Recent molecular evidence, however, has removed the genus Glyphium to the Chaetothyriales in the Eurotiomycetes.[50][51][52] 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.[53] Barr (1979) also originally held a wide view of the Hysteriales,[54] 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.[55] Barr noted (1987) that the relationship of some members of the Hysteriaceae was with the Cucurbitariaceae or with the Pleosporaceae.[4] Finally, Kirk et al. (2001) maintained both the Hysteriaceae and the Mytilinidiaceae in the Hysteriales,[56] but Eriksson (2006) removed the Mytilinidiaceae from the Hysteriales and considered it as Dothideomycetes et 'Chaetothyriomycetes incertae sedis, leaving the Hysteriaceae as the sole family in the Hysteriales.[17]

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.[4] Peridial differences, especially the texture and features related to the peridial base, further separate the two families (Barr 1987).[4] 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.[4] Kutorga and Hawksworth (1997) in their revision of the Patellariaceae did not include Hysteropatella.[57] Initial studies using the nuclear small subunit (nuSSU) rDNA[58][59] have found that Hysterium pulicare and Hysteropatella clavispora (Peck) Seaver formed a clade with high statistical support, distant from other Dothideomycetes, and thus supporting the inclusion of the genus Hysteropatella within the Hysteriaceae.[17]

More recently, Schoch et al. (2006),[60] 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, in agreement with Luttrell (1955).[46] However, it was noted that the hysteriaceous fungi sampled did not form a monophyletic group.[60] 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.[60] 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.[60]

Taken together, classification emphasizing the transitional nature of the hysterothecium, studies in centrum ontogeny 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

[edit]
  • Acrogenospora M.B. Ellis (1971)[61] has 7 species, all of which are anamorphs of genus Farlowiella.[62] the type species is Acrogenospora sphaerocephala (Berk. & Broome) M.B. Ellis (1971).
  • Gloniella Sacc. (1883) contain over 70 species, and the type is Gloniella sardoa Sacc. & Traverso (1883)[63]
  • Gloniopsis De Not. (1847) contains roughly 45 species. The type is Gloniopsis decipiens De Not. (1847).[64]
  • Hysterium Pers. (1797), with roughly 400 species described, is the largest genus of the Hysteriaceae.[65] The type is Hysterium pulicare Ellis.[66]
  • Hysteropatella Rehm (1890) has 8 species,[67] and the type is Hysteropatella prostii (Duby) Rehm (1890).[68]
  • Pseudoscypha J. Reid & Piroz (1966) has one species, the type P. abietis.[69]

References

[edit]
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Other sources

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  • Boehm EWA, Schoch CL, Spatafora JW (2009). On the evolution of the Hysteriaceae and Mytilinidiaceae (Pleosporomycetidae, Dothideomycetes, Ascomycota) using four nuclear genes. Mycol Res 113: 461–479.
  • Boehm EWA, Mugambi GK, Miller AN, Huhndorf SM, Marincowitz S, Spatafora JW, Schoch CL. (2009). A molecular phylogenetic reappraisal of the Hysteriaceae, Mytilinidiaceae, and Gloniaceae (Pleosporomycetidae, Dothideomycetes) with keys to world species. Studies in Mycology 64: 49 – 83.
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