A worldwide nomenclature revision of sequestrate Russula species

Abstract

Before the application of molecular techniques, evolutionary relationships between sequestrate genera and their epigeous counterparts in the Russulaceae were unclear. Based on overwhelming evidence now available, personal observations, and consideration of the International Code for Nomenclature of Algae, Fungi and Plants, we combine the overlapping sequestrate generic names Bucholtzia, Cystangium, Elasmomyces, Gymnomyces, Macowanites, and Martellia with the agaricoid genus Russula. This nomenclatural action follows precedents set by earlier mycologists and continues an effort to create clarity in our understanding of the evolutionary affiliations among sequestrate fungi - particularly the Russulaceae. We also provide the first comprehensive list of described sequestrate species of Russula.

INTRODUCTION

There are many compelling arguments suggesting that sequestrate fruiting habits in fungi resulted from adaptations to abiotic environmental factors and symbiotic associations with vertebrate and invertebrate spore dispersers (Thiers 1984, Cázares & Trappe 1994, Trappe & Claridge 2005, Galante et al. 2011, Gehring et al. 2014). These evolutionary pressures led to the convergence of fruiting macromorphologies in many fungal lineages; however, early mycologists chose to classify fungi with “gasteroid” or “secotioid” fruiting habits in the obsolete class “Gasteromycetes”. At the time, it was thought that this class was distantly related to non-sequestrate (agaricoid) fungi. The evolution of truffle-like fungi from non-sequestrate forms was recognized before the availability of molecular tools (Malençon 1931, Shaffer 1968, Heim 1971, Smith 1973, Trappe 1979, Thiers 1984) and later confirmed (Bruns et al. 1989, Hibbett et al. 1997, Krüger et al. 2001, Peintner et al. 2001, Binder & Bresinsky 2002, Geml 2004, Lebel et al. 2004, Vellinga 2004, Matheny et al. 2006, Albee-Scott 2007, Sheedy et al. 2016).

The spread of molecular approaches in mycology has taken this evolutionary perspective a step further by combining many sequestrate genera with closely affiliated non-sequestrate/gymnocarpic genera. For example, in the Agaricaceae, sequestrate species of Amogaster, Cryptolepiota, Endoptychum, Gyrophragmium, and Longula have been variously described in or transferred to Agaricus, Chlorophyllum, Lepiota, and Macrolepiota (Vellinga 2002, Geml 2004, Geml et al. 2004, Lebel & Syme 2012, Ge & Smith 2013, Lebel 2013, Lebel & Vellinga 2013, Kerrigan 2016). In the Amanitaceae, Torrendia and Amarrendia were combined with Amanita (Justo et al. 2010). In the Bolbitiaceae, the sequestrate genus Descomyces and some members of the genera Setchelliogaster and Timgrovea have been combined with Descolea (Kuhar et al. 2017). In the Boletaceae, a new sequestrate species has been described in Afroboletus, and some Gastroboletus species have been combined with traditionally gymnocarpic genera (Nuhn et al. 2013, Wu et al. 2016, Han 2017). In the Cortinariaceae, Thaxterogaster and some species of Protoglossum have been combined with or described in Cortinarius (Peintner et al. 2002, Danks et al. 2010, Kuhar et al. 2017). In the Entolomataceae, the sequestrate genus Richoniella has been combined with Entoloma (Co-David 2009, Kinoshita et al. 2012). In the Inocybaceae, several sequestrate species have been placed in Inocybe (Braaten et al. 2013). In the Physalacriaceae, members of the sequestrate genus Cribbea have been combined with Oudemansiella (Lebel 2017). In the Psathyrellaceae, the sequestrate Psathyrella secotioides was described (Moreno et al. 2015). In the Strophariaceae, one species of the sequestrate genus Nivatogastrium has been placed in Pholiota and two species of the sequestrate genus Weraroa have been moved to Leratiomyces (Bridge et al. 2008). In the Hymenogastraceae, the sequestrate species Weraroa novae-zelandiae (previously in Strophariaceae) has been recombined and renamed Psilocybe weraroa (Borovička et al. 2011).

The wave of recent nomenclatural changes among sequestrate fungi is analogous to the “one fungus, one name” approach that is in use for revisions of pleomorphic fungus nomenclature (Hawksworth 2015). In the case of sequestrate fungi, it might be better termed “one fungus, several names.” In the next section, we follow the precedents set by these aforementioned mycologists to bring the genus Russula and its associated sequestrate genera up to date with current understandings of sequestrate fungus nomenclature.

SEQUESTRATE SPECIES OF RUSSULA

Since the early 19^th century, new sequestrate Russulae have been placed in one or more of several genera created expressly for them. In early observations of amyloid spore reactions to iodine solutions, Bucholtz (1901, 1903) linked the sequestrate genus Elasmomyces to the Russula lineage and Arcangeliella to Lactarius, even though the phylogenetic affinities were not well understood at the time. The number of sequestrate genera regarded as russuloid in pre-molecular times reached its zenith at five in the seminal paper by Singer & Smith (1960): Cystangium, Elasmomyces, Gymnomyces, Macowanites, and Martellia. A sixth, Bucholtzia, had been validly described by Lohwag (1924) but never accepted by the taxonomist community. Species in two other genera of sequestrate Russulaceae, Arcangeliella and Zelleromyces, are mostly resolved within the epigeous genus Lactarius, although some individual species may eventually be resolved within Russula or other genera in the Russulaceae (Buyck et al. 2008). Each of these 10 genera was circumscribed by characters thought to separate them from the others, but all show various degrees of intergradation (Calonge & Martin 2000, Trappe et al. 2002). Moreover, some sequestrate russuloid species had initially been erroneously assigned to families unrelated to the Russulaceae, including the Agaricaceae, Boletaceae, and Hydnangiaceae. The champion in nomenclatural complexity is Russula krjukowensis, which has been treated in seven genera (see the following species list for the formal new combination).

By the late 20^th century, the phylogenetic relationship of sequestrate russuloid taxa to Russula was firmly demonstrated (Lebel & Trappe 2000, Lebel 2007). Morphological distinctions between some of those genera soon became blurred as additional new species proved to be morphological intermediates. Calonge & Martin (2000), Lebel & Trappe (2000, 2002), and Lebel (2007) attempted to sharpen the generic morphological distinctions, but phylogenetic analyses by Miller et al. (2001, 2006) clearly showed that the Russulaceae are monophyletic, and members of the six named sequestrate russuloid genera are nested among non-sequestrate Russula species with no consistent genetic relationship to each other or to any clade within Russula. The generic names erected to describe sequestrate morphologies of species we now combine with Russula have shown to be nomenclatural artifacts that in hindsight have little bearing on the phylogenetic positions of these taxa. No available phylograms provide evidence that any sequestrate Russula spp. belong to distinct lineages within the genus. The para- or polyphyly within phylogenies that include both sequestrate and non-sequestrate Russula spp. are nomenclatural relics, not genetics.

Over the last twenty years, multiple papers have been published to either describe new sequestrate species in Russula or Lactarius, recombine species from sequestrate genera with Russula or Lactarius, or suggest close or monophyletic relationships between species with the different morphologies (Martín et al. 1999, Miller et al. 2001, Desjardin 2003, Nuytinck et al. 2003, Eberhardt & Verbeken 2004, Lebel & Tonkin 2007, Li et al. 2013, Verbeken et al. 2014, Beenken et al. 2016, Lebel 2017, Looney et al. 2018). It has been widely suggested through published phylogenies that all remaining members of Bucholtzia, Cystangium, Elasmomyces, Gymnomyces, Macowanites, and Martellia should be transferred to Russula. These combinations will reduce the confusion in the current “one fungus, several names” situation and will simplify how we refer to these fungi. Lebel (2007, 2017) did a meticulous job of combining all Australian sequestrate Russulae into Russula. Here we update the remaining described sequestrate Russula species. In an era of genetics, environmental sequencing, and analysis of ecological roles of fungi through sequencing of root tips, having multiple generic names representing fungi that share close genetic affinities leads to further confusion. As stated in the preamble to the 2012 International Code of Nomenclature for Algae, Fungi, and Plants: “The purpose of giving a name to a taxonomic group is not to indicate its characters or history, but to supply a means of referring to it and to indicate its taxonomic rank” (McNeill et al. 2012). Accordingly, we judge that the following nomenclatural revisions of the genus Russula provide the clearest and most accurate information to indicate their taxonomic rank. Some information is lost without the sequestrate generic names; however, this is easily remedied in phylograms by using an arbitrary symbol, colour, or font in association with sequestrate names in a list, caption, text, phylogram, etc., as done by Miller et al. (2001) and Hosaka et al. (2006).

METHODS

We considered genetics whenever possible, but it was impossible to source sequences of type specimens (or new enough specimens) collected from close to type localities to generate a full phylogeny of the species we cover. To resolve the nomenclatural issues in this study, we relied heavily on morphology and reviewed specimens and/or original descriptions of the species we combined to ensure that they belonged in Russula. With increased interest in the sequestrate Russulaceae, we hope that many of these taxa will be re-collected and sequenced to generate a world phylogeny of this group. Genetics would make it possible to resolve the placement of the sequestrate genera Arcangeliella and Zelleromyces within the closely affiliated gymnocarpic genera Lactarius, Lactifluus, and Multifurca.

All specific epithet headings are capitalized and arranged alphabetically in the list below. These species were originally published in the genera Bucholtzia, Cystangium, Elasmomyces, Gymnomyces, Macowanites, or Martellia, as listed in Index Fungorum (plus a few incorrectly placed in non-russuloid genera). In this way, readers can determine how and why an original sequestrate name has been converted to its corresponding Russula name. The protocols for these decisions follow Rossman (2014) and use an amended version of the format provided by Rossman et al. (2016). Sequestrate species with Russula names already or newly published herein are in boldface. The standard nomenclature abbreviation comb. nov. is used when the genus is being combined with Russula but the original specific epithet can still be retained; nom. nov. is used when a new name must be proposed to avoid creating a homonym with a previously described species of Russula. When a new name is needed to avoid producing Russula homonyms, its etymology and other relevant information are noted. MycoBank numbers are assigned and registered to each new combination or new name.

REVISED NOMENCLATURE OF SEQUESTRATE RUSSULA SPP.

Generic type species: Russula emetica (Schaeff.) Pers., Observ. Mycol. (Lipsiae) 1: 100. 1796.

ABIETIS

Russula subabietis Trappe & T.F. Elliott, nom. nov. MycoBank MB821258.

Replaced synonym: Gymnomyces abietis Trappe & Castellano, Mycotaxon 75: 161. 2000, non Russula abietum (J. Blum) Bon, Doc. Mycol. 13 (no. 50): 27. 1983.

Note: subabietis (“almost of the fir”) to avoid confusion between abietis (“of the fir”) with the related epithet abietum (“of firs”).

ACRIS

Russula acerba Trappe & T.F. Elliott,nom. nov. MycoBank MB821259.

Replaced synonym: Macowanites acris Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 86. 1960, non Russula acris Steinhaus, Hedwigia 27: 51. 1888.

Etymology and note: acerba (“bitter”) to avoid producing a homonym with the earlier acris (“sharp”) of Russula acris Steinhaus (1888).

AGARICINUS

Russula agaricina (Kalchbr. ex Berk.) Trappe & T.F. Elliott,comb. nov. MycoBank MB 825252.

Basionym: Macowania agaricina Kalchbr. ex Berk., Gard. Chron. 5: 785. 1876.

Synonyms: Macowanites agaricinus (Kalchbr. ex Berk.) Kalchbr., Grevillea 10(55): 107. 1882. Russula agaricina (Kalchbr. ex Berk.) T. Lebel, [as ‘(Kalchbr.) T. Lebel.’], Austral. Syst. Bot. 20: 360. 2007, nom. inval. (Art. 41.5).

Note: Macowinia agaricina is a legitimate name although published under Kalchbrenner’s illegitimate genus name Macowania (Art. 55.1) and has to be considered in terms of priority issues (Art. 11.4). Lebel (in Lebel & Tonkin 2007) intended to reallocate this name to Russula but did not introduce a valid combination (basionym not cited). She only referred to Macowanites agaricinus, which is only an indirect reference to the true basionym and not sufficient to validate the combination (Art. 41.5).

ALBA

Russula stevemilleri Trappe & T.F. Elliott,nom. nov. MycoBank MB821262.

Replaced synonym: Hydnangium album Harkn., Proc. Calif. Acad. Sci., Ser. 3, Bot. 1: 251. 1899, non Russula alba Velen., České Houby 1: 149. 1920.

Synonyms: Martellia alba (Harkn.) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 41. 1961.

Gymnomyces albus (Harkn.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 198. 2002.

Etymology and note: In honor of Dr. Steve Miller for his continuing research on taxonomy and phylogeny of sequestrate and other Russula species. This new name is erected to avoid producing a homonym with Russula alba Velen. (1920).

ALBIDIGLEBA

Russula albidigleba (Singer & A.H. Sm.) Trappe & T.F. Elliott, comb. nov. MycoBank MB821263.

Basionym: Macowanites albidiglebus Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 85. 1960.

ALBIDOFLAVA

Russula albidoflava T. Lebel, Austral. Syst. Bot. 20: 361. 2007. MycoBank MB532175.

ALBOBRUNNEA

Russula albobrunnea T. Lebel, Austral. Syst. Bot. 20: 362. 2007. MycoBank MB532176.

ALPINUS

Russula alpicola Trappe & T.F. Elliott, nom. nov. MycoBank MB821264.

Replaced synonym: Macowanites alpinus Zeller, Mycologia 39: 291. 1947, non Russula alpina (A. Blytt & Rostr.) F.H. Møller & Jul. Schäff., Ann. Mycol. 38(2/4): 333. 1940.

Synonym: Elasmomyces alpinus (Zeller) Zeller, Mycologia 40: 643. 1948.

Etymology and note: alpicola (“dweller of alps”), to avoid producing a homonym with the earlier Russula alpina (A. Blytt & Rostr.) F.H. Møller & Jul. Schäff. (1940).

AMMOPHILUS

Russula ammophila (J.M. Vidal & Calonge) Trappe & T.F. Elliott,comb. nov. MycoBank MB821265.

Basionym: Gymnomyces ammophilus J.M. Vidal & Calonge, Bol. Soc. Micol. Madrid 24: 66. 1999.

Synonym: Macowanites ammophilus (J.M. Vidal & Calonge) J.M. Vidal & Calonge, Revista Catal. Micol. 24: 70. 2002.

ARENICOLA

Russula arenicola (S.L. Mill. & D. Mitch.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821266.

Basionym: Macowanites arenicola S.L. Mill. & D. Mitch., Mycotaxon 89: 284. 2004.

BALPINEUM

Russula balpinea (Grgur.) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820015.

Basionym: Cystangium balpineum Grgur., Larger Fungi of South Australia (Adelaide): 53 (1997).

BISPORUM

Russula bispora (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820016.

Basionym: Cystangium bisporum T. Lebel, Austral. Syst. Bot. 16: 373. 2003.

BORANUPENSIS

Russula boranupensis (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820018.

Basionym: Gymnomyces boranupensis T. Lebel, Austral. Syst. Bot. 16: 403. 2003.

BRUNNEONIGRA

Russula brunneonigra T. Lebel, Austral. Syst. Bot. 20: 363. 2007. MycoBank MB532177.

BRUNNESCENS

Russula shafferi Trappe & T.F. Elliott,nom. nov. MycoBank MB821312.

Replaced synonym: Martellia brunnescens Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 37. 1961, non Russula brunnescens Murrill, Lloydia 8: 264. 1946 (1945).

Synonym: Gymnomyces brunnescens (Singer & A.H. Sm.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002.

Etymology and note: To honor Dr. Robert Shaffer for his important contributions to knowledge of North American Russulae. This new name is erected to avoid producing a homonym with the earlier Russula brunnescens Murrill (1945).

CALIFORNICA

Russula californica (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821336.

Basionym: Martellia californica Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 33. 1961.

Synonym: Gymnomyces californicus (Singer & A.H. Sm.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002, non Russula californiensis Burl., Mycologia 28: 262. 1936.

CANDIDUS

Note: Originally described as Hydnangium candidum Tul. & C. Tul. in 1843, this species was recombined by Zeller & Dodge (1935) as Sclerogaster candidus (Tul. & C. Tul.) Zeller & C.W. Dodge, and then by Vidal (2005) as Macowanites candidus (Tul. & C. Tul.) J.M. Vidal. According to Kirk (2010), S. candidus in the Sclerogastraceae, not the Russulaceae, is the correct name.

CAPITIS-ORAE

Russula capitis-orae (Dring) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820020.

Basionym: Elasmomyces capitis-orae Dring, in Dring & Pegler, Kew Bull. 32(3): 564. 1978.

Synonym: Cystangium capitis-orae (Dring) T. Lebel, Mycotaxon 81: 197. 2002.

CHLORINOSMUS

Russula chlorineolens Trappe & T.F. Elliott,nom. nov. MycoBank MB821355.

Replaced synonym: Macowanites chlorinosmus A.H. Sm. & Trappe, Mycologia 55: 423. 1963, non Russula chlorinosma Burl., Mycologia 16: 22. 1924.

Etymology and note: “chlorineolens” (smelling like chlorine), to avoid producing a homonym with the earlier, similarly smelling Russula chlorinosmus Burl. (1924).

CINNAMOMEA

Russula unicalifornica Trappe & T.F. Elliott,nom. nov. MycoBank MB821338.

Replaced synonym: Gymnomyces cinnamomeus Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 51. 1960, non Russula cinnamomea Banning, Bot. Gaz. 6(1): 166. 1881.

Etymology and note: “unicalifornica” contraction of “University of California” in reference to the site of the type collection to avoid producing a homonym with the earlier Russula cinnamomea Banning (1881).

CITRINA

Russula harknessii Trappe & T.F. Elliott,nom. nov. MycoBank MB821339.

Replaced synonym: Octaviania citrina Harkn., Proc. Calif. Acad. Sci., Ser. 3, Bot. 1: 252. 1899, non Russula citrina Gillet, Revue mycol., Toulouse 3(no. 11): 5. 1881, non Macowanites citrinus Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 69. 1960.

Synonyms: Hydnangium citrinum (Harkn.) Zeller & C.W. Dodge, Ann. Missouri. Bot. Gard. 22: 371. 1935.

Gymnomyces citrinus (Harkn.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002.

Etymology and note: In honor of H. W. Harkness, pioneer in taxonomy of North American sequestrate fungi. This new name is erected to avoid producing a homonym with the earlier Russula citrina Gillet (1881).

CITRINUS

Russula alleinstanleyae T.F. Elliott & Trappe,nom. nov. MycoBank MB823193.

Replaced synonym: Macowanites citrinus Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 69. 1960, non Russula citrina Gillet, Revue Mycol., (Toulouse) 3(no. 11): 5 (1881), non Gymnomyces citrinus (Harkn.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002.

Etymology and note: In honor of mycologist and educator Allein Stanley for her more than 40 years of dedicated service to the North American Mycological Association. This new name is erected to avoid producing a homonym with the earlier Russula citrina Gillet (1881).

CLAVATUM

Russula clavata (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820021.

Basionym: Cystangium clavatum T. Lebel, Austral. Syst. Bot. 16: 375. 2003.

CLELANDII

Russula aurantirosea T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820058.

Replaced synonym: Gymnomyces clelandii T. Lebel, Austral. Syst. Bot. 16: 404. 2003, non Russula clelandii O.K. Mill. & R.N. Hilton, Sydowia 39: 128. 1987.

COMPACTA

Russula mattsmithii Trappe & T.F. Elliott, nom. nov. MycoBank MB821354.

Replaced synonym: Gymnomyces compactus Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 53. 1960, non Russula compacta Frost, Rep. (Annual) New York State Mus. Nat. Hist. 32: 32. 1879.

Etymology and note: In honor of Dr. Matt Smith for his extensive research on taxonomy and ecology of sequestrate fungi. This new name is erected to avoid producing a homonym with the earlier Russula compacta Frost (1879).

COSTATISPORUS

Russula costatispora (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820022.

Basionym: Gymnomyces costatisporus T. Lebel, Austral. Syst. Bot. 16: 405. 2003.CREMEA

Russula dodgei Trappe & T.F. Elliott,nom. nov. MycoBank MB824950.

Replaced synonym: Arcangeliella cremea Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 22: 367. 1935, non Russula cremea (Murrill) Singer, Lilloa 22: 711. 1951.

Synonyms: Martellia cremea (Zeller & C.W. Dodge) Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 45. 1961 (1960).

Gymnomyces cremeus (Zeller & C.W. Dodge) Trappe et al., Mycotaxon 81: 199. 2002.

Etymology and note: In honor of C.W. Dodge, who collaborated with S. M. Zeller in describing numerous new species of sequestrate fungi, including this taxon and to avoid confusion with Russula cremea (Murrill) Singer.

CRISTATA

Russula korystospora T. Lebel, Muelleria 36: 12. 2017. MycoBank MB821530.

Replaced synonym: Gymnomyces cristatus T. Lebel, New Zealand J. Bot. 40: 491. 2002, non Russula cristata Romagn., Bull. Mens. Soc. Linn. Soc. Bot. Lyon 31(1): 177. 1962.

DEPAUPERATUM

Russula depauperata (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821344.

Basionym: Cystangium depauperatum Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 69. 1960.

DOMINGUEZIAE

Russula laurae Trappe & T.F. Elliott, nom. nov. MycoBank MB821357.

Replaced synonym: Cystangium domingueziae Nouhra & Trierv.-Per., Mycologia 107: 94. 2014, non Gymnomyces dominguezii Mor.-Arr., J. Gómez & Calonge, Mycol. Res. 103: 215. 1999.

Etymology and note: In honor of Argentine mycologist, Laura S. Dominguez after whom the species was originally named. This new name is erected to avoid producing a homonym with the earlier and similar name Gymnomyces dominguezii Mor.-Arr., J. Gómez & Calonge (see following entry).

DOMINGUEZII

Russula dominguezii (Mor.-Arrl, J. Gómez & Calonge) Trappe & T.F. Elliott,comb. nov. MycoBank MB821346.

Basionym: Gymnomyces dominguezii Mor.-Arr., J. Gómez & Calonge, Mycol. Res. 103: 215. 1999.

DURANGENSIS

Russula durangensis (Guzmán) Trappe & T.F. Elliott,comb. nov. MycoBank MB821358.

Basionym: Macowanites durangensis Guzmán, Revista Mex. Micol. 4: 118. 1988.

EBURNEA

Russula eburnea (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820023.

Basionym: Gymnomyces eburneus T. Lebel, Austral. Syst. Bot. 16: 408. 2003.

ECHINOSPORUS

Russula tombrunsii Trappe & T.F. Elliott,nom. nov. MycoBank MB823196.

Replaced synonym: Macowanites echinosporus Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 6: 57. 1919, non Russula echinospora Singer, Bull. Soc. Mycol. France 55: 270. 1940, non Russula echinospora R. Heim, Rev. Mycol. (Toulouse) 32: 207. 1967, non Cystangium echinosporum (Zeller & C.W. Dodge) Trappe, T. Lebel & Castellano, Mycotaxon 81: 197. 2002, non Elasmomyces echinosporus Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 22: 370. 1935.

Etymology and note: In honor of Dr. Tom Bruns for his career-long research on taxonomy, phylogeny and ecology of ectomycorrhizal fungi. This new name is erected to avoid producing a homonym of R. echinospora Singer (1940).

EILDONENSIS

Russula eildonensis (G.W. Beaton et al.) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820024

Basionym: Gymnomyces eildonensis G.W. Beaton et al., Kew Bull. 39: 680. 1984.

ELLIPSOSPORUM

Russula ellipsospora (Zeller) Trappe & T.F. Elliott,comb. nov. MycoBank MB821361.

Basionym: Hydnangium ellipsosporum Zeller, Mycologia 31: 13. 1939.

Synonyms: Martellia ellipsospora (Zeller) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 30. 1960.

Gymnomyces ellipsosporus (Zeller) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002.

FALLAX

Russula similaris Trappe & T.F. Elliott,nom. nov. MycoBank MB821362.

Replaced synonym: Martellia fallax Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 34. 1960, non Russula fallax (Schaeff.) Fr., Hymenomyc. Eur. (Upsaliae): 449. 1874.

Synonym: Gymnomyces fallax (Singer & A.H. Sm.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002.

Etymology and note: “similaris” in reference to the similarity of the two epithets cited above, “fallax” (deceptive) deceiving their respective authors, and to avoid producing a homonym with the earlier R. fallax (Schaef.) Fr. (1874).

FERRUGINASCENS

Russula ferruginascens (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821363.

Basionym: Gymnomyces ferruginascens Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 54. 1960.

FLAVOVIRENS

Russula olivaceoflava T. Lebel, Muelleria 36: 12. 2017. MycoBank MB821532.

Replaced synonym: Cystangium flavovirens T. Lebel, Austral. Syst. Bot. 16: 377. 2003, non Russula flavovirens J. Bommer & M. Rousseau, Bull. Soc. Roy. Bot. Belgique 23(1): 310. 1884.

FOETENS

Russula taetra Trappe & T.F. Elliott,nom. nov. MycoBank MB821365.

Replaced synonym: Martellia foetens Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 36. 1960, non Russula foetens Pers., Observ. Mycol. (Lipsiae) 1: 102. 1796.

Synonym: Gymnomyces foetens (Singer & A.H. Sm.) Trappe et al., Mycotaxon 81: 199. 2002.

Etymology and note: taetra (“foul, offensive”), a related term to express the original epithet foetens (“stinking”) to avoid producing a homonym of the earlier Russula foetens Pers. (1796).

FRAGRANS

Russula aromatica Trappe & T.F. Elliott,nom. nov. MycoBank MB821366.

Replaced synonym: Martellia fragrans A.H. Sm., Mycologia 55: 437. 1963.

Synonym: Gymnomyces fragrans (A.H. Sm.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002, non Russula fragrans Romagn., Bull. Mens. Soc. Linn. Soc. Bot. Lyon 23: 112. 1954.

Etymology and note: “aromatica”, a related term to express the original epithet fragrans “becoming fragrant” to avoid producing a homonym of the earlier Russula fragrans Romagn. (1954).

FULVESCENS

Russula vilgalysii Trappe & T.F. Elliott,nom. nov. MycoBank MB821367.

Replaced synonym: Macowanites fulvescens Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 84. 1960, non Russula fulvescens Burl., N. Amer. Fl. (New York) 9: 229. 1915.

Etymology and note: In honor of Dr. Rytas Vilgalys for his research and teaching of all aspects of mycology. This new name is erected to avoid producing a homonym of the earlier Russula fulvescens Burl. (1915).

FULVISPORA

Russula fulvispora (A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821368.

Basionym: Martellia fulvispora A.H. Sm., Mycologia 55: 438. 1963.

Synonym: Gymnomyces fulvisporus (A.H. Sm.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 199. 2002.

FURCATISPINA

Russula furcatispina (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820025.

Basionym: Gymnomyces furcatispinus T. Lebel, Austral. Syst. Bot. 16: 411. 2003.

FUSCOVIOLACEUS

Russula fuscoviolaceus (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821371.

Basionym: Macowanites fuscoviolaceus Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 87. 1960.

FUSCUS

Russula spinispora T. Lebel, Muelleria 36: 12. 2017. MycoBank MB821534.

Replaced synonym: Gymnomyces fuscus T. Lebel, New Zealand J. Bot. 40: 495. 2002, non Russula fusca Quél., Compt. Rend. Assoc. Franç. Avancem. Sci. 15: 486. 1887.

GALBANA

Russula galbana T. Lebel, Austral. Syst. Bot. 20: 365. 2007. MycoBank MB532179.

GALILEENSIS

Russula galileensis (M.M. Moser, Binyam. & Aviz.-Hersh.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821373.

Basionym: Macowanites galileensis M.M. Moser, Binyam. & Aviz.-Hersh., Trans. Brit. Mycol. Soc. 68: 371. 1977.

GAMUNDIAE

Russula gamundiae (Nouhra & Trierv.-Per.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821374.

Basionym: Cystangium gamundiae Nouhra & Trierv.-Per., Mycologia 107: 95. 2014.

GILKEYAE

Russula gilkeyae (Zeller & C.W. Dodge) Trappe & T.F. Elliott,comb. nov. MycoBank MB821375.

Basionym: Hydnangium gilkeyae Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 22: 371. 1935.

Synonyms: Octaviania gilkeyae (Zeller & C.W. Dodge) Svrček, Fl. ČSR, B-1, Gasteromycetes: 192. 1958.

Martellia gilkeyae (Zeller & C.W. Dodge) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 43. 1960.

Gymnomyces gilkeyae (Zeller & C.W. Dodge) Trappe et al., Mycotaxon 81: 199. 2002.

GLAREA

Russula glarea (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820026.

Basionym: Gymnomyces glarea T. Lebel, Austral. Syst. Bot. 16: 412. 2003.

GRANDIHYPHATUM

Russula grandihyphata (Nouhra & Trierv.-Per.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821377.

Basionym: Cystangium grandihyphatum Nouhra & Trierv.-Per., Mycologia 107: 96. 2014.

IDAHOENSIS

Russula idahoensis (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821378.

Basionym: Martellia idahoensis Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 43. 1960.

Synonym: Cystangium idahoense (Singer & A.H. Sm.) Trappe et al. [as ‘idahoensis’], Mycotaxon 81: 197. 2002.

ILICIS

Russula vidalii Trappe & T.F. Elliott,nom. nov. MycoBank MB821379.

Replaced synonym: Gymnomyces ilicis J.M. Vidal & Llistos., Rivista Micol. 38: 160. 1995, non Russula ilicis Romagn. et al., Bull. Trimestriel Soc. Mycol. France 88: 33. 1972.

Etymology and note: In honor of J.D. Vidal, Spanish Catalonian mycologist who has specialized in taxonomy of sequestrate fungi. This new name is erected to avoid producing a homonym with the earlier Russula ilicis Romagn. (1972).

IODIOLENS

Russula iodiolens (A.H. Sm. & V.L. Wells) Trappe & T.F. Elliott,comb. nov. MycoBank MB821380.

Basionym: Macowanites iodiolens, A.H. Sm. & V.L. Wells, Mycologia 55: 425. 1963.

JAVANICA

Russula boedijnii Trappe & T.F. Elliott,nom. nov. MycoBank MB823199.

Replaced synonym: Hydnangium javanicum Henn., Hedwigia 40(Beibl.): 27. 1901, non Russula javanica Sacc. & P. Syd., Syll. Fung. (Abellini) 16: 46. 1902.

Synonym: Gymnomyces javanicus (Henn.) M.E. Sm. & Schmull, Mycol. Progr. 10: 253. 2011.

Etymology and note: In honor of Dutch mycologist K.B. Boedijn, who specialized in taxonomy of Indonesian fungi including hypogeous species. This new name is erected to avoid producing a homonym with the earlier Russula javanica Sacc. & P. Syd. (1902).

KERMESINA

Russula kermesina T. Lebel, Austral. Syst. Bot. 20: 377. 2007. MycoBank MB531311.

Replaced synonym: Macowanites carmineus McNabb, New Zealand J. Bot. 9: 359. 1971, non Russula carminea (Jul. Schäff.) Kühner & Romagn., in Romagnesi, Russules d’Europe Afr. Nord, Essai sur la Valeur Taxinomique et Specifique des Characteres des Spores et des Revetements: 447. 1967.

KRJUKOWENSIS

Russula krjukowensis (Bucholtz) Trappe & T.F. Elliott,comb. nov. MycoBank MB823514.

Basionym: Secotium krjukowense Bucholtz, Hedwigia 40: 314. 1901.

Synonyms: Secotium michailowskianum Bucholtz [as ‘michailowskjanum’], Hedwigia 40: 315. 1901.

Elasmomyces michailowskianus (Bucholtz) Sacc. & D. Sacc. [as ‘michailowskjanus’], Syll. Fung. (Abellini) 17: 218. 1905.

Elasmomyces krjukowensis f. pleurotopsis Bucholtz, Bull. Soc. Imp. Naturalistes Moscou, ser. 2, 24: 463. 1907.

Bucholtzia krjukowensis (Bucholtz) Lohwag, Oesterr. Bot. Z. 73(7-9): 173. 1924.

Arcangeliella krjukowensis var. michailowskiana (Bucholtz) Zeller & C.W. Dodge, Ann. Missouri. Bot. Gard. 22: 368. 1935.

Arcangeliella krjukowensis (Bucholtz) Zeller & C.W. Dodge, Ann. Mo. Bot. Gdn 22: 368. 1935. var. krjukowensis.

Hydnangium krjukowense (Bucholtz) Svrček, Fl. ČSR, B-1, Gasteromycetes: 206. 1958. var. krjukowense.

Macowanites krjukowensis (Bucholtz) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 77. 1960.

LEUCOCARPUS

Russula leucocarpa (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820035.

Basionym: Gymnomyces leucocarpus T. Lebel, New Zealand J. Bot. 40: 497. 2002.

LONGISPORUS

Russula longispora (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820036.

Basionym: Gymnomyces longisporus T. Lebel, Austral. Syst. Bot. 16: 415. 2003.

LONGISTERIGMATUM

Russula longisterigmata (Nouhra & Trierv.-Per.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821384.

Basionym: Cystangium longisterigmatum Nouhra & Trierv.-Per., Mycologia 107: 97. 2014.

LUTEIROSEA

Russula luteirosea (Bougher) T. Lebel, Muelleria 36: 13 (2017). MycoBank MB822048.

Basionym: Macowanites luteiroseus Bougher, Mycotaxon 63: 38. 1997.

Synonym: Russula luteirosea (Bougher) T. Lebel, Austral. Syst. Bot. 20(4): 378. 2007, nom. inval.

LUTEOBRUNNEUM

Russula luteobrunnea (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820037.

Basionym: Cystangium luteobrunneum T. Lebel, Austral. Syst. Bot. 16: 380. 2003.

LUTEOLUS

Russula luteola (Harkn.) Trappe & T.F. Elliott, comb. nov. MycoBank MB821386.

Basionym: Hydnangium luteolum Harkn., Proc. Calif. Acad. Sci., Ser. 3, Bot. 1: 251. 1899.

Synonym: Gymnomyces luteolus (Harkn.) Trappe et al., Mycotaxon 81: 200. 2002, non Macowanites luteolus A.H. Sm. & Trappe, Mycologia 55: 427. 1963.

LUTEOLA

Russula stricklandorum T.F. Elliott & Trappe,nom. nov. MycoBank MB821387.

Replaced synonym: Macowanites luteolus A.H. Sm. & Trappe, Mycologia 55: 427. 1963, non Russula luteola (Harkn.) Trappe & T.F. Elliott (this paper, see preceding entry).

Etymology and note: stricklandorum, in honor of Bob and Babs Strickland for their support of the natural sciences and contributions to conservation. This new name is erected to avoid producing a homonym with the preceding Russula luteola (Harkn.) Trappe & T.F. Elliott (2017).

LYMANENSE

Russula lymanensis (Cázares & Trappe) Trappe & T.F. Elliott,comb. nov. MycoBank MB821388.

Basionym: Macowanites lymanensis Cázares & Trappe, Mycotaxon 42: 335. 1991.

Synonym: Cystangium lymanense (Cázares & Trappe) Trappe et al. [as ‘lymanensis’], Mycotaxon 81: 197. 2002.

MACROCYSTIDIUM

Russula macrocystidia (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820038.

Basionym: Cystangium macrocystidium T. Lebel, Austral. Syst. Bot. 16: 380. 2003.

MACULATA

Russula extramaculata Trappe & T.F. Elliott, nom. nov. MycoBank MB821390.

Replaced synonym: Martellia maculata Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 28. [1960] 1961, non Russula maculata Quél., Bull. Soc. Bot. France 24: 323. [1877] 1878.

Synonym: Cystangium maculatum (Singer & A.H. Sm.) Trappe et al., Mycotaxon 81: 197. 2002.

Etymology and note: extramaculatum (“outside maculata”) to avoid producing a homonym with the earlier Russula maculata Quél. (1878).

MATTIROLOANUS

Russula mattiroloana (Cavara) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820039.

Basionym: Elasmomyces mattiroloanus Cavara [as ‘mattirolianus’], Malpighia 11(9–10): 426. 1897.

Synonyms: Secotium mattiroloanum (Cavara) E. Fisch., in Engler & Prantl, Nat. Pflanzenfam., Teil. I (Leipzig) 1: 301. 1900.

Macowanites mattiroloanus (Cavara) T. Lebel & Trappe (as ‘mattirolianus’), Mycologia 92: 1194. 2000.

MEDITERRANEA

Russula mediterranea (G. Moreno et al.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821392.

Basionym: Martellia mediterranea G. Moreno et al., Mycotaxon 42: 227. 1991.

MEDLOCKII

Russula medlockii (Trappe & Castellano) Trappe & T.F. Elliott,comb. nov. MycoBank MB821393.

Basionym: Martellia medlockii Trappe & Castellano, Mycologia 78: 918. 1986.

Synonym: Cystangium medlockii (Trappe & Castellano) Trappe et al., Mycotaxon 81: 197. 2002.

MEGASPORUS

Russula megaspora (Rodway) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820040.

Basionym: Gymnomyces megasporus Rodway, Pap. & Proc. Roy. Soc. Tasmania ‘1925’: 168. 1926 (1925).

Synonyms: Octaviania megaspora (Rodway) G. Cunn., New Zealand J. Sci. Technol. 22: 300B. 1941.

Cystangium megasporum (Rodway) T. Lebel & Castellano, Mycologia 94: 335. 2002.

MESSAPICOIDES

Russula messapicoides (Llistos. & J.M. Vidal) Trappe & T.F. Elliott,comb. nov. MycoBank MB821395.

Basionym: Macowanites messapicoides Llistos. & J.M. Vidal, Rivista Micol. 38: 155. 1995.

MEXICANUS

Russula guzmanii Trappe & T.F. Elliott,nom. nov. MycoBank MB821396.

Replaced synonym: Macowanites mexicanus Guzmán, Revista Mex. Micol. 4: 116. 1988, non Russula mexicana Burl., Mycologia 3: 26. 1911.

Etymology and note: In honor of Prof. Gastón Guzmán, much accomplished Mexican mycologist and taxonomist. This new name is erected to avoid producing a homonym of the earlier Russula mexicana Burl. (1911).

MISTIFORMIS

Russula mistiformis (Mattir.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821397.

Basionym: Martellia mistiformis Mattir., Malpighia 14: 78. 1900.

Synonyms: Hydnangium mistiforme (Mattir.) Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 22: 372. 1935.

Octaviania mistiformis (Mattir.) Svrček, Fl. ČSR, B-1, Gasteromycetes: 192. 1958.

Gymnomyces mistiformis (Mattir.) T. Lebel & Trappe, Mycologia 92: 1999. 2000.

MOLLIS

Russula castellanoi Trappe & T.F. Elliott,nom. nov. MycoBank MB821398.

Replaced synonym: Macowanites mollis Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 79. 1960, non Russula mollis Quél., Compt. Rend. Assoc. Franç. Avancem. Sci. 11: 397. 1883 (1882).

Synonym: Elasmomyces mollis (Singer & A.H. Sm.) Pegler & T.W.K. Young, Trans. Brit. Mycol. Soc. 72: 368. 1979.

Etymology and note: In honor of Dr. Michael (Caz) Castellano for his career-long fascination with and research on sequestrate fungi. This new name is erected to avoid producing a homonym with the earlier Russula mollis Quél. (1882).

MONOSPORA

Russula monospora (Boud. & Pat.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821399.

Basionym: Hydnangium monosporum Boud. & Pat., J. Bot., Paris 2: 445. 1888.

Synonyms: Octaviania monospora (Boud. & Pat.) Lloyd, Mycol. Writ. 7(Letter 67): 1141. 1922.

Martellia monospora (Boud. & Pat.) Astier & Pacioni, Doc. Mycol. 28(nos 109–110): 9. 1998.

MONOSPORUS

Russula stewartii Trappe & T.F. Elliott,nom. nov. MycoBank MB821400.

Replaced synonym: Gymnomyces monosporus E.L. Stewart & Trappe, Mycotaxon 2: 209. 1975.

Etymology and note: In honor of Dr. Elwin L. Stewart, mycologist and plant pathologist, for his contributions to taxonomy of sequestrate fungi. This new name is erected to avoid producing a homonym of the earlier Hydnangium monosporum Boud. & Pat. (1888). See previous entry.

MONTICOLA

Russula monticola (Harkn.) Trappe & T.F. Elliott,comb. nov. MycoBank MB821401.

Basionym: Octaviania monticola Harkn., Proc. Calif. Acad. Sci., Ser. 3, Bot. 1: 254. 1899.

Synonyms: Hydnangium monticola (Harkn.) Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 22: 372. 1935.

Martellia monticola (Harkn.) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 40. 1961 (1960).

Gymnomyces monticola (Harkn.) Trappe et al., Mycotaxon 81: 200. 2002.

NANJINGENSIS

Russula nanjingensis (B. Liu & K. Tao) Trappe & T.F. Elliott,comb. nov. MycoBank MB823202.

Basionym: Martellia nanjingensis B. Liu & K. Tao, Acta Mycol. Sin. 12: 103. 1993.

Synonyms: Gymnomyces nanjingensis (B. Liu & K. Tao) Trappe et al., Mycotaxon 81: 200. 2002.

Arcangeliella nanjingensis (B. Liu & K. Tao) J.M. Vidal, Revista Catal. Micol. 26: 73. 2005 (2004).

NAUSEOSUS

Russula hunsuckeri T.F. Elliott & Trappe,nom. nov. MycoBank MB823515.

Replaced synonym: Macowanites nauseosus A.H. Sm., Mycologia 55: 428. 1963, non Russula nauseosa (Pers.) Fr., Epicr. Syst. Mycol. (Upsaliae): 363. [1836–1838] 1838.

Etymology and note: In honor of naturalist, mycologist, botanist, and philosopher Dr. Robert Hunsucker. This new name is erected to avoid producing a homonym with the earlier Russula nauseosa (Pers.) Fr. (1838).

NONDISTINCTA

Russulanondistincta (Trappe & Castellano) Trappe & T.F. Elliott,comb. nov. MycoBank MB823207.

Basionym: Gymnomyces nondistincta Trappe & Castellano, Mycotaxon 75: 166. 2000.

NOTHOFAGI

Russula nothofagi (E. Horak) Trappe & T.F. Elliott,comb. nov. MycoBank MB823517.

Basionym: Elasmomyces nothofagi E. Horak, Sydowia 17(1–6): 211. [1963] 1962.

Synonym: Cystangium nothofagi (E. Horak) Trappe et al., Mycotaxon 81: 198. 2002.

OCCIDENTALIS

Russula thiersii Trappe & T.F. Elliott,nom. nov. MycoBank MB823208.

Replaced synonym: Martellia occidentalis Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 42. 1960, non Russula occidentalis (Singer) Singer, Lilloa 22: 705. [1949] 1951, non Russula occidentalis Bon, Doc. Mycol. 12(no. 46): 32. 1982, nom. illeg.

Etymology and note: In honor of Dr. Harry Thiers for his long interest in and taxonomic studies of fungi and the Russulaceae. This new name is erected to avoid producing a homonym with the earlier Russula occidentalis Singer (1951).

ODORATUS

Russula odorifera Trappe & T.F. Elliott,nom. nov. MycoBank MB823209.

Replaced synonym: Elasmomyces odoratus Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 57. 1960, non Russula odorata Romagn., Bull. Mens. Soc. Linn. Soc. Bot. Lyon 19: 76. 1950.

Synonym: Macowanites odoratus (Singer & A.H. Sm.) Trappe et al., Mycotaxon 81: 202. 2002.

Etymology and note: odorifera (bearing an odor), to avoid producing a homonym with the earlier Russula odorata Romagn. (1950).

OLIDUS

Russula olida (A.H. Sm.) Trappe & T.F. Elliott, comb. nov. MycoBank MB823210.

Basionym: Macowanites olidus A.H. Sm., Mycologia 55: 428. 1963.

OREGONENSIS

Russula oregonensis (Zeller) Trappe & T.F. Elliott,comb. nov. MycoBank MB823211.

Basionym: Hydnangium oregonense Zeller, Mycologia 33: 200. 1941

Synonyms: Martellia oregonensis (Zeller) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 27. 1961 (1960).

Cystangium oregonense (Zeller) Trappe et al., Mycotaxon 81: 198. 2002.

PALLIDUS

Russula paneeroides T. Lebel, Muelleria 36: 12. 2017. MycoBank MB821533.

Replaced synonym: Gymnomyces pallidus Massee & Rodway, in Massee, Bull. Misc. Inform. Kew: 125. 1898, non Russula pallida P. Karst., Hedwigia 35: 43. 1896.

Synonym: Octaviania pallida (Massee & Rodway) G. Cunn., Proc. Linn. Soc. New South Wales 60: 119. 1935.

PARKSII

Russula parksii (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823213.

Basionym: Gymnomyces parksii Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 55. 1960.

PARVISAXOIDES

Russula parvisaxoides (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB822047.

Basionym: Gymnomyces parvisaxoides T. Lebel, New Zealand J. Bot. 40: 498. 2002.

PHYMATODISPORUM

Russula phymatodispora (G.W. Beaton et al.) T. Lebel Muelleria 36: 12. 2017. MycoBank MB820041.

Basionym: Cystangium phymatodisporum G.W. Beaton et al., Kew Bull. 39: 672. 1984.

PILA

Russula pila (Pat.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823216.

Basionym: Hydnangium pila Pat., Bull. Soc. Mycol. France 26: 202. 1910.

Synonyms: Octaviania pila (Pat.) Svrček, Fl. ČSR, B-1, Gasteromycetes: 199. 1958.

Martellia pila (Pat.) J.M. Vidal, Bull. Soc. Catalana Micol. 14–15: 172. 1991.

Gymnomyces pila (Pat.) Trappe, T. Lebel & Castellano, Mycotaxon 81: 200. 2002.

PILOSUS

Russula pilosa (Zeller & C.W. Dodge) Trappe & T.F. Elliott,comb. nov. MycoBank MB823217.

Basionym: Arcangeliella pilosa Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 22: 368. 1935.

Synonyms: Elasmomyces pilosus (Zeller & C.W. Dodge) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 66. 1960.

Macowanites pilosus (Zeller & C.W. Dodge) Trappe et al., Mycotaxon 81: 202. 2002.

PILOSELLA

Russula pilosella T. Lebel, Austral. Syst. Bot. 20: 379. 2007. MycoBank MB532181.Replaced synonym: Hydnangium tomentosum J.W. Cribb, Pap. Dept. Bot. Univ. Queensland 3: 251. 1958, non Russula tomentosa Buyck, Bull. Jard. Bot. Natl. Belg. 58: 476. 1988.

Synonyms: Martellia tomentosa (J.W. Cribb) A.H. Sm., Mycologia 54: 631. 1963.

PINETI

Russula pineti (Singer) Trappe & T.F. Elliott,comb. nov. MycoBank MB823218.

Basionym: Cystangium pineti Singer, Mycol. Helv. 1: 417. 1985.

PINICOLA

Russula piniamans Trappe & T.F. Elliott,nom. nov. MycoBank MB823225.

Replaced synonym: Macowanites pinicola A.H. Sm., Mycologia 55: 430. 1963, non Russula pinicola Murrill, Lloydia 6: 213. 1943.

Etymology and note: piniamans (“friend of pine”), to avoid producing a homonym of the earlier Russula pinicola Murrill (1943)

PISIGLAREA

Russula pisiglarea (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820042.

Basionym: Cystangium pisiglarea T. Lebel, Austral. Syst. Bot. 16: 383. 2003.

POLYCHROMUM

Russula collubrina T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820059.

Replaced synonym: Cystangium polychromum Trappe & Claridge, Australas. Mycol. 22: 33. 2003, non Russula polychroma Singer ex Hora, Trans. Brit. Mycol. Soc. 43: 457. 1960.

PSEUDOEMETICUS

Russula nancyweberae Trappe & T.F. Elliott,nom. nov. MycoBank MB823545.

Replaced synonym: Macowanites pseudoemeticus A.H. Sm., Mycologia 55: 431. 1963, non Russula pseudoemetica Secr. ex Singer, Hedwigia 66: 190. 1926.

Etymology and note: In honor of Dr. Nancy Weber, for her extensive research on taxonomy of epigeous Ascomycetes, especially the genus Morchella. This new name is erected to avoid producing a homonym with the earlier Russula pseudoemetica Secr. ex Singer (1926).

PTEROSPERMUS

Russula pterosperma (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820043.

Basionym: Gymnomyces pterospermus T. Lebel, Austral. Syst. Bot. 16: 416. 2003.

PUMICOIDEA

Russula pumicoidea T. Lebel, Austral. Syst. Bot. 20: 367. 2007. MycoBank MB532182.

REDDELLII

Russula reddellii T. Lebel, Austral. Syst. Bot. 20: 369. 2007. MycoBank MB532183.

REDOLENS

Russula osphranticarpa T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820061.

Replaced synonym: Octaviania redolens G. Cunn., New Zealand J. Sci. Technol. 23B: 172. 1942, non Russula redolens Burl., Mycologia 13: 133. 1921.

Synonym: Gymnomyces redolens (G. Cunn.) Pfister, Occas. Papers Farlow Herb. Cryptog. Bot. 9: 43. 1976.

RODWAYI

Russula leonardii T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820060.

Replaced synonym: Gymnomyces rodwayi T. Lebel, Austral. Syst. Bot. 16: 418. 2003, non Russula rodwayi (Massee) T. Lebel, Muelleria 36: 11. 2017.

Russula rodwayi (Massee) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820044.

Basionym: Secotium rodwayi Massee, Bull. Misc. Inform. Kew: 158. 1901.

Synonyms: Elasmomyces rodwayi (Massee) Zeller, Mycologia 40: 642. 1948.

Cystangium rodwayi (Massee) A.H. Sm., Mycologia 54: 633. 1963.

ROGERSII

Russula rogersii (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823546.

Basionym: Octavianina rogersii Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 12. 1960.

Synonym: Gymnomyces rogersii (Singer & A.H. Sm.) Trappe et al. Mycotaxon 81: 201. 2002.

ROLFALEXII

Russula rolfalexii (Trappe et al.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823552.

Basionym: Gymnomyces rolfalexii Trappe et al., Mycotaxon 81: 201. 2002.

Synonym: Martellia parksii Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 32. 1960, non Gymnomyces parksii Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 55. 1960. See Russula parksii (Singer & A.H. Sm.) Trappe & T.F. Elliott this publication.

ROSEIPES

Russula spataforae Trappe & T.F. Elliott,nom. nov. MycoBank MB823547.

Replaced synonym: Elasmomyces roseipes Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 64. 1960, non Russula roseipes Secr. ex Bres., Fung. Trident. 1: 37. 1881.

Synonym: Macowanites roseipes (Singer & A.H. Sm.) Pegler & T.W.K. Young, Trans. Brit. Mycol. Soc. 72: 363. 1979.

Etymology and note: In honor of Dr. Joey Spatafora for his teaching and extensive research on phylogeny and taxonomy of macrofungi. This new name is established to avoid producing a homonym with Russula roseipes Secr. ex Bres. (1881).

ROSEOMACULATUS

Russula roseomaculata (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823548.

Basionym: Gymnomyces roseomaculatus Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 50. 1960.

ROSTRATICYSTIDIA

Russula rostraticystidia T. Lebel, Austral. Syst. Bot. 20: 371. 2007. MycoBank MB532184.

RUBROLUTEA

Russula rubrolutea (T. Lebel) T. Lebel, Muelleria 36: 14. 2017. MycoBank MB822049.

Basionym: Macowanites rubroluteus T. Lebel, New Zealand J. Bot. 40: 505. 2002.

Synonym: Russula rubrolutea (T. Lebel) T. Lebel, Austral. Syst. Bot. 20: 378. 2007, nom. inval.

RUSSULOIDES

Russula russuloides (Setch.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823549.

Basionym: Elasmomyces russuloides Setch., J. Mycol. 13: 240. 1907.

Synonym: Macowanites russuloides (Setch.) Trappe et al., Mycotaxon 81: 202. 2002.

SICHUANENSIS

Russula sichuanensis G.J. Li & H.A. Wen, Mycotaxon 124: 179. 2013 MycoBank MB804645.

SEMINUDUS

Russula seminuda (Massee & Rodway) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820045.

Basionym: Gymnomyces seminudus Massee & Rodway, Bull. Misc. Inform. Kew: 125. 1898.

Synonyms: Arcangeliella seminuda (Massee & Rodway) Zeller & C.W. Dodge, Ann. Missouri Bot. Gard. 23: 617. 1936.

Octaviania seminuda (Massee & Rodway) G. Cunn., Trans. & Proc. Roy. Soc. New Zealand 67: 408. 1938.

Cystangium seminudum (Massee & Rodway) T. Lebel & Castellano, Mycologia 94: 337. 2002.

SESSILE

Russula sessilis (Massee & Rodway) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820046.

Basionym: Secotium sessile Massee & Rodway, Pap. & Proc. Roy. Soc. Tasmania ‘1911’: 31. [1911] 1912 [var. sessile].

Synonyms: Elasmomyces sessilis (Massee & Rodway) Rodway, Pap. & Proc. Roy. Soc. Tasmania ‘1924’: 8. [1924] 1925.

Cystangium sessile (Massee & Rodway) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 27. 1960.

SETCHELLIANUS

Russula setchelliana (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823550.

Basionym: Macowanites setchellianus Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 92. 1960.

SETIGER

Russula setigera (Zeller) Trappe & T.F. Elliott,comb. nov. MycoBank MB823551.

Basionym: Hydnangium setigerum Zeller, Mycologia 31: 14. 1939.

Synonyms: Martellia setigera (Zeller) Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 39. 1960.

Gymnomyces setiger (Zeller) Trappe et al. [as ‚setigerus‘], Mycotaxon 81: 201. 2002.SHULTZIAE

Russula shultziae (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820047.

Basionym: Cystangium shultziae T. Lebel, Austral. Syst. Bot. 16: 385. 2003.

SICHUANENSIS

Russula sichuanensis G.J. Li & H.A. Wen, Mycotaxon 124: 179. 2013. MycoBank MB804645.

SINUATA

Russula sinuata T. Lebel, Austral. Syst. Bot. 20: 374. 2007. MycoBank MB532185.

SOLIDUS

Russula solida (Rodway) Trappe & T.F. Elliott, comb. nov. MycoBank MB823522.

Basionym: Gymnomyces solidus Rodway, Pap. & Proc. Roy. Soc. Tasmania ‘1921’: 157. 1921.

SPARSUM

Russula sparsa (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820048.

Basionym: Cystangium sparsum T. Lebel, Austral. Syst. Bot. 16: 388. 2003.

STIPITATUS

Russula stipitata (H.A. Peters) Trappe & T.F. Elliott, comb. nov. MycoBank MB823523.

Basionym: Elasmomyces stipitatus H.A. Peters, Mycologia 54: 112. 1962.

Synonym: Macowanites stipitatus (H.A. Peters) Trappe et al., Mycotaxon 81: 202. 2002.

SUBALPINUS

Russula orsonmilleri Trappe & T.F. Elliott,nom. nov. MycoBank MB823524.

Replaced synonym: Martellia subalpina A.H. Sm., Mycologia 55: 439. 1963, non Russula subalpina O.K. Mill., Arctic and Alpine Mycology, First International Symposium on Arcto-Alpine Mycology, 1980 (Seattle): 136. 1982.

Synonym: Gymnomyces subalpinus (A.H. Sm.) Trappe et al., Mycotaxon 81: 201. 2002.

Etymology and note: In honor of Dr. Orson K. Miller Jr. for his wide ranging contributions to taxonomy of North American fungi, including sequestrate and subalpine species. This new name is erected to avoid producing a homonym with the earlier Russula subalpina O.K. Miller (1963).

SUBFULVUS

Russula subfulva (Singer & A.H. Sm.) Trappe & T.F. Elliott, comb. nov. MycoBank MB823525.

Basionym: Martellia subfulva Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 35. 1960.

Synonym: Gymnomyces subfulvus (Singer & A.H. Sm.) Trappe et al., Mycotaxon 81: 201. 2002.

SUBLEVISPORUS

Russula andaluciana T.F. Elliott & Trappe, nom. nov. MycoBank MB823526.

Replaced synonym: Gymnomyces sublevisporus Mor.-Arr. et al., Revista Catal. Micol. 24: 179. 2002, non Russula sublevispora (Romagn.) Kühner & Romagn., in Romagnesi, Russules d’Europe Afr. Nord, Essai sur la Valeur Taxinomique et Specifique des Characteres des Spores et des Revetements: 299. 1967.

Etymology and note: In reference to Andalucia, the region where it was originally collected, and to avoid producing a homonym with Russula sublevispora (Romagn.) Kühner & Romagn (1967).

SUBOCHRACEUS

Russula subochracea (A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823527.

Basionym: Martellia subochracea A.H. Sm., Mycologia 55: 440. 1963.

Synonym: Gymnomyces subochraceus (A.H. Sm.) Trappe et al., Mycotaxon 81: 201. 2002.

SUBOLIVACEUS

Russula subolivacea (A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823528.

Basionym: Macowanites subolivaceus A.H. Sm., Mycologia 55: 432. 1963.

SUBROSACEUS

Russula subrosacea (A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823529.

Basionym: Macowanites subrosaceus A.H. Sm., Mycologia 55: 433. 1963.

TAPAWERA

Russula tapawera (T. Lebel) T. Lebel, Austral. Syst. Bot. 20: 379. 2007. MycoBank MB531354.

Basionym: Macowanites tapawera T. Lebel, New Zealand J. Bot. 40: 507. 2002.

THAXTERI

Russula thaxteri (Singer) Trappe & T.F. Elliott,comb. nov. MycoBank MB823530.

Basionym: Martellia thaxteri Singer, Beih. Nova Hedwigia 29: 357. 1969.

Synonym: Cystangium thaxteri (Singer) Trappe et al., Mycotaxon 81: 198. 2002.

THEODOROUI

Russula theodoroui (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820049.

Basionym: Cystangium theodoroui T. Lebel, Austral. Syst. Bot. 16: 390. 2003.

TRAPPEI

Russula trappei (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820050.Basionym: Cystangium trappei T. Lebel, Austral. Syst. Bot. 16: 392. 2003.

VARIABILISPORA

Russula variabilispora (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823531.

Basionym: Martellia variabilispora Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 27. 1960.

Synonym: Cystangium variabilisporum (Singer & A.H. Sm.) Trappe et al., Mycotaxon 81: 198. 2002.

VARIISPORA

Russula variispora T. Lebel, Austral. Syst. Bot. 20: 375. 2007. MycoBank MB532186.

VESICULOSA

Russula marshallorum T.F. Elliott & Trappe,nom. nov. MycoBank MB823553.

Replaced synonym: Martellia vesiculosa Singer & A.H. Sm., Mem. Torrey Bot. Club 21: 29. 1960, non Russula vesciculosa (Coker & Couch) Trappe & T.F. Elliott (this paper, see below).

Synonym: Cystangium vesiculosum (Singer & A.H. Sm.) Trappe et al., Mycotaxon 81: 198. 2002.

Etymology and note: In honor of Australian truffle-growing Marshall family for their strong support of Australasian truffle research and conservation. This new name is erected to avoid producing a homonym with Russula vesciculosa (see following entry).

VESICULOSUS

Russula vesiculosa (Coker & Couch) Trappe & T.F. Elliott,comb. nov. MycoBank MB823554.

Basionym: Gymnomyces vesiculosus Coker & Couch, Gasteromycetes of the Eastern U.S. and Canada (Chapel Hill): 23. 1928.

Synonym: Hydnangium vesiculosum (Coker & Couch) Zeller, Mycologia 40: 641. 1948.

VINACEODORUS

Russula vinaceodora (Calonge & J.M. Vidal) Trappe & T.F. Elliott,comb. nov. MycoBank MB823532.

Basionym: Macowanites vinaceodorus Calonge & J.M. Vidal, Mycotaxon 79: 2. 2001.

VINICOLOR

Russula vinicolor (A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823533.

Basionym: Macowanites vinicolor A.H. Sm., Mycologia 55: 434. 1963.

WESTRESII

Russula westresii (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820051.

Basionym: Gymnomyces westresii T. Lebel, Austral. Syst. Bot. 16: 420. 2003.

WIRRABARENSIS

Russula wirrabarensis (Grgur.) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820052.

Basionym: Gymnomyces wirrabarensis Grgur., Larger Fungi of South Australia (Adelaide): 86. 1997.

XANTHOCARPUM

Russula xanthocarpa (T. Lebel) T. Lebel, Muelleria 36: 12. 2017. MycoBank MB820053.

Basionym: Cystangium xanthocarpum T. Lebel, Austral. Syst. Bot. 16: 394. 2003.

XANTHOSPORUS

Russula xanthospora (Hawker) Trappe & T.F. Elliott. comb. nov. MycoBank MB823534.

Basionym: Hydnangium carneum var. xanthosporum Hawker, Trans. Brit. Mycol. Soc. 35: 281. 1952.

Synonym: Gymnomyces xanthosporus (Hawker) A.H. Sm., Mycologia 54: 635. 1963.

XEROPHILUS

Russula xerophila (M.E. Sm. & Trappe) Trappe & T.F. Elliott, comb. nov. MycoBank MB823535.

Basionym: Gymnomyces xerophilus M.E. Sm. & Trappe, Mycol. Res. 110: 577. 2006.

YUNNANENSIS

Russula zangii Trappe & T.F. Elliott,nom. nov. MycoBank MB823536.

Replaced synonym: Macowanites yunnanensis M. Zang, Acta Bot. Yunnan. 21: 37. 1999, non Russula yunnanensis (Singer) Singer, Mycologia 34: 72. 1942.

Etymology and note: In honor of M. Zang, original describer of the species. This new name is erected to avoid producing a homonym with the earlier Russula yunnanensis (Singer) Singer (1942).

ZELLERIANUS

Russula zelleriana (Singer & A.H. Sm.) Trappe & T.F. Elliott,comb. nov. MycoBank MB823537.

Basionym: Elasmomyces zellerianus Singer & A.H. Sm., Mem. Torrey Bot. Club 21(3): 61. 1960.

Synonym: Macowanites zellerianus (Singer & A.H. Sm.) Trappe et al., Mycotaxon 81: 203. 2002.