Tremella

Tremella
	Gelatinous fruit body of Tremella mesenterica
Scientific classification
Domain:	Eukaryota
Kingdom:	Fungi
Division:	Basidiomycota
Class:	Tremellomycetes
Order:	Tremellales
Family:	Tremellaceae
Genus:	Tremella; Pers. (1801)
Type species
	Tremella mesenterica; (Schaeff.) Retz.
Synonyms
	Epidochium Fr.; Gyraria Nees; Nakaiomyces Kobayasi;

Tremella is a genus of fungi in the family Tremellaceae. All Tremella species are parasites of other fungi and most produce anamorphic yeast states. Basidiocarps (fruit bodies), when produced, are gelatinous and are colloquially classed among the "jelly fungi". Over 100 species of Tremella (in its wide sense) are currently recognized worldwide. One species, Tremella fuciformis, is commercially cultivated for food.

Taxonomy

History

Tremella was one of the original genera created by Linnaeus in his Species Plantarum of 1753. The name comes from the Latin tremere meaning "to tremble".[1] Linnaeus placed Tremella in the algae, including within it a variety of gelatinous growths, including seaweeds, cyanobacteria, and myxomycetes, as well as fungi. Subsequent authors added additional species to this mix, until Persoon revised Tremella in 1794 and 1801, repositioning the genus within the fungi.[2]

Persoon's reinterpretation of Tremella was sufficiently radical to be considered a separate genus (Tremella Pers.) from that originally created by Linnaeus (Tremella L.).[2] Tremella Pers. has now been conserved under the International Code of Nomenclature for algae, fungi, and plants, with Tremella mesenterica as the type species.[3]

Current status

Molecular research, based on cladistic analysis of DNA sequences, has shown that Tremella (as previously understood) is polyphyletic (and hence artificial), with most species not closely related to the type.[4][5][6][7] Accordingly, some species have been transferred to new genera and new families: Tremella foliacea and related species are now placed in the genus Phaeotremella within the family Phaeotremellaceae; Tremella encephala and related species are now placed in the genus Naematelia within the Naemateliaceae; Tremella moriformis and related species are now placed in the genus Pseudotremella within the Bulleraceae; and Tremella polyporina is now placed in the genus Carcinomyces within the Carcinomycetaceae.[8] Several other species groups have not yet been renamed, pending further research.[8]

More than 500 species have been described in Tremella, but most of these are old names of doubtful application or old names for species later transferred to other genera. In its strict sense, the genus Tremella now contains some 30-40 species, including the type Tremella mesenterica and the cultivated species T. fuciformis.[9]

Description

Fruit bodies (when present) are gelatinous. In some species they are small (under 5 mm across) and pustular to pulvinate (cushion-shaped). In others they are much larger (up to 150 mm across) and may be variously lobed, cephaliform (like a brain, with folds and ridges), or foliose (with leaf-like or seaweed-like fronds). Many Tremella species, however, are hymenial parasites, producing spores within the fruit bodies of their hosts, and are only visible microscopically.[4]

Microscopic characters

Tremella species produce hyphae that are typically (but not always) clamped and have haustorial cells from which hyphal filaments seek out and penetrate the hyphae of the host.[10] The basidia are "tremelloid" (globose to ellipsoid, sometimes stalked, and vertically or diagonally septate), giving rise to long, sinuous sterigmata or epibasidia on which the basidiospores are produced. These spores are smooth, globose to ellipsoid, and germinate by hyphal tube or by yeast cells. Conidiophores are often present, producing conidiospores that are similar to yeast cells.[4]

Habitat and distribution

Species are mainly parasitic on wood-rotting fungi in the phyla Ascomycota and Basidiomycota,[11] particularly on species that occur on dead attached branches. Hosts include members of the corticioid fungi and Dacrymycetales in the Basidiomycota and species of Diaporthe, other Sordariomycetes, and lichens in the Ascomycota. Some Tremella species parasitize the fruit bodies of their hosts, others parasitize the mycelium within the wood.

As a group, Tremella species occur worldwide, though individual species may have a more restricted distribution.

Species and hosts

The list below includes species of Tremella (in the wide sense) that have recently been described or redescribed based on fruit bodies. Species based on yeasts are not included. Some additional older species may also be valid, but lack a modern description. The type locality (but not the wider distribution) is given for each species together with the host fungus, where known. Species belonging to Tremella in the strict sense are marked as such, as are those that have been transferred to new genera.

Name	Current Status	Type Locality	Host
Tremella anaptychiae		Spain	Anaptychia ciliaris[12]
Tremella anomala		Brazil	Hypoxylon spp[13]
Tremella arachispora		Cameroon	unknown[14]
Tremella armeniaca	Tremella sensu stricto[9]	Costa Rica	Xylaria sp?[15]
Tremella aurantia	Naematelia aurantia	USA	Stereum hirsutum[4][16]
Tremella aurantialba	Naematelia aurantialba	China	Stereum hirsutum[17]
Tremella aurantiolutea		Mexico	unknown[18]
Tremella australe	Tremella sensu stricto[19]	China	unknown[19]
Tremella australiensis		Australia	Stereum spp[4]
Tremella boninensis	Tremella sensu stricto[9]	Japan	unknown[20]
Tremella brasiliensis	Tremella sensu stricto[9]	Brazil	unknown[4][16][21]
Tremella callunicola		Scotland	Aleurodiscus norvegicus[22]
Tremella caloceraticola		Denmark	Calocera cornea[23]
Tremella caloplacae		Greece	Caloplaca spp[24](as Tremella sp. 1)
Tremella candelariellae		Luxembourg	Candelariella spp[24]
Tremella cephalodiicola		Papua New Guinea	Psoroma pannarioides[24]
Tremella cerebriformis	Tremella sensu stricto[9]	Taiwan	unknown[4]
Tremella cetrariicola		Scotland	Cetraria spp[24]
Tremella cheejenii	Tremella sensu stricto[9]	China	unknown[9]
Tremella christiansenii		Denmark	Physcia spp[24]
Tremella cladoniae		Germany	Cladonia spp[24]
Tremella clavisterigma		Guatemala	unknown[25]
Tremella coalescens	Tremella sensu stricto[8]	USA	unknown[4]
Tremella coccocarpiae		Philippines	Coccocarpia rottleri[24]
Tremella coffeicolor		Bermuda	unknown[13][16](as T. auricularia)
Tremella colpomaticola		Denmark	Colpoma quercinum[23]
Tremella compacta		Brazil	unknown[26]
Tremella coppinsii		Sarawak	Platismatia spp[24]
Tremella dendrographae		USA	Dendrographa minor[24]
Tremella dactylobasidia		Spain	Dendrothele macrosporae[27]
Tremella discicola		Belgium	Mollisia and Pyrenopeziza spp[28]
Tremella dumontii		Ecuador	unknown[18]
Tremella durissima		Argentina	unknown[25]
Tremella dysenterica	Tremella sensu stricto[9]	Brazil	unknown[13][16]
Tremella encephala	Naematelia encephala	Europe	Stereum sanguinolentum[4][29]
Tremella erythrina	Tremella sensu stricto[9]	China	unknown[9]
Tremella everniae		China	Evernia mesomorpha[24]
Tremella exigua		France	Diaporthe spp[4]
Tremella fibulifera	Tremella sensu stricto[9]	Brazil	unknown
Tremella fimbriata	Phaeotremella fimbriata	Europe	Stereum rugosum
Tremella flammea	Tremella sensu stricto[9]	Japan
Tremella flava	Tremella sensu stricto[9]	Taiwan	Hypoxylon spp[4]
Tremella foliacea	Phaeotremella foliacea	Europe	Stereum sanguinolentum
Tremella frondosa	Phaeotremella frondosa	Europe	Stereum spp
Tremella fuciformis	Tremella sensu stricto[9]	Brazil	Annulohypoxylon archeri[30][31] and other Annulohypoxylon and/or Hypoxylon spp.[4][16][31]
Tremella fungicola		Denmark	Mollisia cinerea[32]
Tremella fuscosuccinea	Phaeotremella fuscosuccinea	Taiwan	unknown[4]
Tremella giraffa		Germany	Dacrymyces spp[4][33]
Tremella globispora	Tremella sensu stricto[9]	England	Diaporthe spp[4]
Tremella graphidastrae		Papua New Guinea	Graphidastra multiformis[24]
Tremella guangxiensis	Tremella sensu stricto[19]	China	unknown[19]
Tremella guttiformis		Sri Lanka	unknown[34]
Tremella haematommatis		USA	Haematomma puniceum[24]
Tremella hainanensis	Tremella sensu stricto[9]	China	unknown[35]
Tremella harrisii		USA	Polymeridium catapastum[24]
Tremella huuskonenii			Raesaenenia huuskonenii[36]
Tremella hymenophaga		Spain	Scytinostroma odoratum[37]
Tremella hypocenomycis		Finland	Hypocenomyce spp[24][38]
Tremella hypogymniae		France	Hypogymnia spp[24]
Tremella iduensis	Tremella sensu stricto[9]	Japan	Hypoxylon spp [20]
Tremella indecorata		Norway	Diatrype bullata[38]
Tremella invasa		Denmark	Trechispora spp[22]
Tremella karstenii		Finland	Colpoma juniperi[23][38]
Tremella latispora	Tremella sensu stricto[19]	China	unknown[19]
Tremella leptogii		Peru	Leptogium sp.[24]
Tremella lethariae		Canada	Letharia vulpina[39]
Tremella lichenicola		Luxembourg	Mycoblastus fucatus[24]
Tremella lilacea		Costa Rica	Diaporthe sp?[15]
Tremella lloydiae-candidae	Tremella sensu stricto[9]	Japan	unknown
Tremella lobariacearum		Madeira	Lobariaceae spp[24]
Tremella macroceratis		Norway	Cladonia macroceras[24]
Tremella mangensis		China	unknown[35]
Tremella mesenterella	Tremella sensu stricto[9]	Canada	Peniophora spp[21]
Tremella mesenterica	Tremella sensu stricto[9]	Sweden	Peniophora spp[21]
Tremella microcarpa		Papua New Guinea	lichen sp.[24]
Tremella microspora	Naematelia microspora	South Africa	Stereum spp[4]
Tremella monospora		Peru	Leptogium sp.[24]
Tremella montis-wilhelmii		Papua New Guinea	Normandina simodense sp.[24]
Tremella moriformis	Pseudotremella moriformis	England	Diaporthe spp[4]
Tremella mycetophiloides	Phaeotremella mycetophiloides	Japan	Aleurodiscus amorphus and A. grantii[40]
Tremella mycophaga	Phaeotremella mycophaga	Canada	Aleurodiscus amorphus and A. grantii[40]
Tremella nashii		USA	Usnea sorediifera[41]
Tremella nephromatis		Canada	Nephroma parile[24]
Tremella nieblae		USA	Niebla cephalota[41]
Tremella nigrifacta		Costa Rica	Diatrypella sp.[15]
Tremella nivalis	Pseudotremella nivalis	Taiwan	Diatrype spp[4]
Tremella normandinae		Hawaii	Normandina pulchella[24]
Tremella obscura		USA	Dacrymyces spp[33]
Tremella occultifuroidea		Taiwan	Dacrymyces spp[42]
Tremella olens	Tremella sensu stricto[19]	Australia	unknown[43]
Tremella papuana		Papua New Guinea	Hypogymnia pseudobitteriana[24]
Tremella parmeliarum		Papua New Guinea	Parmotrema spp[24]
Tremella parmeliellae		Papua New Guinea	Parmeliella foliicola[24]
Tremella penetrans		Denmark	Dacrymyces spp[33]
Tremella pertusariae		Northern Ireland	Pertusaria spp[24]
Tremella phaeographidis		England	Phaeographis spp[24]
Tremella phaeographinae		USA	Phaeographina spp[24]
Tremella phaeophysciae		Denmark	Phaeophyscia orbicularis[24]
Tremella philippinensis	Tremella sensu stricto[9]	Philippines	unknown
Tremella poilkavensis		India	Biscogniauxia sp.
Tremella polyporina	Carcinomyces polyporinus	Scotland	Postia spp[33]
Tremella protoparmeliae		England	Protoparmelia spp[24]
Tremella pseudocyphellariae		Mauritius	Pseudocyphellaria desfontainii
Tremella psoroglaenae		Papua New Guinea	Psoroglaena spp[24]
Tremella psoromicola		Chile	Psoroma spp[24]
Tremella puncteliae		Mexico	Punctelia borreri
Tremella puncteliotegens		New Zealand	Punctelia borreri
Tremella purpurascentis		USA	Dirinaria purpurascens
Tremella pyrenaica		France	Lecanora polytropa
Tremella ramalinae		Mexico	Ramalina spp[24]
Tremella ramboidiae		USA	Ramboldia haematites
Tremella resupinata	Tremella sensu stricto[9]	Taiwan	Hypoxylon spp[4]
Tremella rhabdodisci		Japan	Rhabdodiscus inalbescens
Tremella rhizocarpicola		Norway
Tremella rinodinae		Denmark	Rinodina spp[24]
Tremella robusta		USA	Roccellina fransciscana
Tremella rosea		Austria	unknown
Tremella roseolutescens	Tremella sensu stricto[9]	Costa Rica	unknown[15]
Tremella rubromaculata	Tremella sensu stricto[9]	Guatemala	unknown[25]
Tremella salmonea	Tremella sensu stricto[9]	China	unknown[9]
Tremella samoensis	Tremella sensu stricto[9]	Samoa	unknown
Tremella sanguinea	Phaeotremella[44]	China	Stereum sp.
Tremella santessonii		Zimbabwe	Usnea spp[24]
Tremella sarcographae		Brazil	Sarcographa medusulina
Tremella sarniensis		Guernsey	Phanerochaete sordida[22]
Tremella seclusa		France	unknown
Tremella silvae-dravedae		Denmark	unknown[23]
Tremella simplex	Phaeotremella simplex	Canada	Aleurodiscus amorphus[22][40]
Tremella spicifera		Belgium	Massarina arundinacea[45]
Tremella steidleri		Czech Republic	Stereum hirsutum[29]
Tremella stevensiana		Australia	Usnea confusa[24]
Tremella stictae		Rwanda	Sticta weigelii[24]
Tremella strigulae		USA	Strigula stigmatella
Tremella subalpina		Russia	unknown
Tremella subencephala		Canada	Acanthophysium lividocoeruleum[40]
Tremella subfibulifera	Tremella sensu stricto[19]	Brazil	unknown[19]
Tremella subrubiginosa	Tremella sensu stricto[9]	Panama	unknown
Tremella sulcariae		China	Sulcaria sulcata[24]
Tremella synarthoniae		USA	Synarthonia inconspicua
Tremella taiwanensis	Tremella sensu stricto[9]	Taiwan	unknown[4]
Tremella tawa	Tremella sensu stricto[9]	New Zealand	unknown[46]
Tremella telleriae		Spain	Postia spp[37]
Tremella teloschistis		USA	Teloschistes exilis
Tremella tornabeae		Canary Islands	Tornabea scutellifera
Tremella translucens		Scotland	Lophodermium spp[47](as Sirotrema translucens)
Tremella tremelloides	Naematelia[34]	USA	Stereum sp.[34]
Tremella tropica	Tremella sensu stricto[9]	Taiwan	unknown[4]
Tremella tubulosae		Scotland	Hypogymnia tubulosa
Tremella tuckerae		USA	Ramalina sinensis[41]
Tremella umbilicariae		Peru	Umbilicaria sp.
Tremella vasifera		Germany	unknown[4]
Tremella versicolor		England	Peniophora spp[22][40]
Tremella vesiculosa		New Zealand	Xylaria sp?[46]
Tremella volcanagua		Guatemala	unknown[25]
Tremella wedinii		USA
Tremella wirthii		Germany	Protoparmelia sp.[24]
Tremella wrightii		Cuba	unknown[16]
Tremella xanthomendozae		USA	Xanthomendoza weberi
Tremella yokohamensis	Tremella sensu stricto[9]	Japan	unknown
Tremella zamorae		USA	Lecanora louisianae
Tremella zhejiangensis	Tremella sensu stricto[48]	China	unknown[48]

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[Rea1922-1] Rea C. (1922). 'British Basidiomycetaceae. A handbook of the larger British fungi. Cambridge: Cambridge University Press. p. 799.

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