Fungal Diversity Three type specimens designated in

Petersen, R.H.1*, Desjardin, D.E.2 and Krüger, D.3

1Ecology and Evolutionary Biology, University of Tennessee, Knoxville, TN 37996-1100 USA 2Department of Biology, San Francisco State University, San Francisco, CA 94132 3Helmholtz-Zentrum für Umweltforschung GmbH-UFZ, Department of Soil Ecology, Theodor-Lieser-Str. 4, D-06120 Halle/Saale, Germany

Petersen, R.H., Desjardin, D.E. and Krüger, D. (2008). Three type specimens designated in Oudemansiella. Fungal Diversity 32: 81-96.

Type specimens are designated for three species epithets in Oudemansiella: O. canarii, an apparently pantropical species, O. platensis, type species of the genus and apparently distributed in tropical and subtropical New World, and O. mucida, common and indigenous on Fagus wood in Europe. Detailed descriptions are furnished for the designated types of all three and the backgrounds of all three epithets are traced.

Key words: nomenclature, type specimens

Article Information Received 30 March 2008 Accepted 21 May 2008 Published online 30 September 2008 *Corresponding author: R. Petersen; e-mail: [email protected]

Introduction Von Höhnel made several attempts to understand the complex of taxa which he Farr (1973: 924-925) furnished an abbre- thought surrounded Oudemansiella. At first viated summary of literature which, if followed, (1908. Frag. Mykol. V, Sitzungber. Kaiserl. provides a recapitulation of confusion concern- Akad. Wissenschaft. Wien 117: 985-1031), in ing the distribution and of Spegaz- reviewing Hennings’ genus Phaeolimacium zini’s (1880) species, Oudemansia [nom. illeg., [which von Höhnel considered to be the same later renamed as Oudemansiella; Spegazzini, as Oudemansiella, and which he recapitulated 1881 ] platensis. later (1911. Resultate der Revision von Paul Saccardo (1887: 653-654) considered that Hennings’ Pilzgattungen. Ann. Mycol. 9: 166- Spegazzini’s Agaricus (Tricho-loma) platensis 175.)], von Höhnel listed five species he (Fungi Argent. pug. 1: 161. 1880) was an considered to belong in Oudemansiella: 1) Ag. earlier introduction, probably based only on the () apalosarcus Berk. & Broome; 2) specific epithet. Ag. (Collybia) magisterium Berk. & Broome; Hennings (1900, in Engler & Prantl, 3) Ag. (Collybia) euphyllus Berk & Broome; 4) Natürl. Pflanzenf. 1 (Abt. 1): 221) gave the Phaeolimacium bulbosum Henn.; and 5) distribution of O. platensis as Argentina and Pluteus macrosporus Henn.. After lengthy Ecuador. discussion, von Höhnel (1908: 1007) emended Rick (1905: 238) listed the species for the characters of Spegazzini’s Oude-mansiella Rio Grande do Sul in southern Brazil, and and listed two species, O. platensis Speg. and discussed characters which seem diagnostic for O. apalosarca (Berk. & Broome) Höhn., as Oudemansiella (i.e. pleurocystidia too large for accepted in Oudemansiella, with the Mycena), although not necessarily for O. aforementioned five names as synonyms (but platensis. If correctly identified, distribution not clearly synonyms of what). The following was enlarged to include Brazilian subtropics, year (von Höhnel. 1909. Frag. Mykol. 6, although he considered basidiomata as repre- Sitzungber. Kaiserl. Akad. Wissenschaft. Wien senting a species of Mycena parasitized by a 118: 276) he recombined Junghühn’s Agaricus “Phycomycete.” canarii as . In a further 81 attempt, von Höhnel (1910. Frag. Mykol. 12, that at least two species, macroscopically Sitzungber. Kaiserl. Akad. Wissenschaft. Wien similar but differing microscopically, were 119: 877-958) listed three species [O. mucida involved. He concluded that O. platensis, (Schrad.) Höhn., O. cheimonophyllum (Berk. & characterized by a polycystoderm pilepellis, M.A.Curtis) Höhn., O. canarii (Jungh.) Höhn.] was a New World species, while O. canarii, each with its own synonyms. Previously (von exhibiting a disarticulated trichoderm, was Höhnel 1908) in prose, Armillaria mucida was common in the western Pacific and Southeast asso-ciated although it was characterized by the Asia. If these names (O. canarii, O. platensis) presence of a well-developed annulus at represented different organisms, there was maturity, but later (1913. Frag. Mykol. XV, reason to deduce that one organism might be Sitzungber. Kaiserl. Akad. Wissenschaft. Wien Old World and the other New World. A 122: 255-309), emphasizing basidiome solution to this problem lies in the microana- development and the role of the universal veil, tomy of their type specimens. he conjectured that A. mucida was better in Horak (1968: 425) did not report on the Armillaria or Lepiota, where annulate veils type specimen of O. platensis for his redescrip- were prominent. tion of the species, although one of the consulted Petch (1910) took up what he called specimens [Uruguay, Depto. Montevideo, III- Oudemansiella apalosarca (Berk. & Broome) IV.1935, leg. Herter, Plantae Uruguayenses Höhn., listing as synonyms the same names as Exs. (G)] was collected across the estuary of those reported by von Höhnel in 1911 (see the Rio de la Plata from the topotype location. above). Petch’s informal description states: “It Horak wrote: “... Diese pantropische Art inter- [O. apalosarca] resembles an Armillaria, but pretieren wir nach Singer (1950: 185; 1964: lacks a ring, and the stalk is distinct from the 145) …” The second consulted specimen pileus. Its general diaphanous appearance (Matheson Hammock, X.1942, leg & det R recalls A. mucida. Young specimens (5 mm. Singer as O. canarii; FH) was collected in diameter) are usually dark brown, studded with Florida, USA, and was not distinguished by minute white fragments of the universal veil.” Horak from the Uruguay collection. The white floccules are recognized as common In order to understand the species of in the modern concept of O. canarii and of O. Oudemansiella, it is necessary to test Corner’s platensis. Previously, Petch (1907: 33-34) cited hypothesis against the traditional (Singerian) examples of difficulties in interpretations of extensive synonymy. It was soon discovered, Ceylon fungi, using Collybia magisterium however, that Spegazzini’s type specimen of O. Berk. & Broome as an example of a species platensis was worthless for detailed study (as with conflicting descriptions and at least three previously reported by Baroni and Ortiz, 2002), names. Parenthetically, Petch (1910) also and that no Junghühn type specimen survived transferred Marasmius subaurantiacus Berk. & for Agaricus canarii. The intent of this paper, Broome to Oudemansiella. therefore, is to designate and describe type With such a plethora of names and specimens for these two organisms. We also disparate reports of geographic sightings, it is designate a neotype specimen for the common little wonder that Singer (1964) and Pegler European species, O. mucida. In the case of O. (1986) could list a full page of synonyms under mucida, a detailed description is necessary only O. canarii (including multiple species epithets). to secure the type specimen to detailed taxono- Furthermore, O. canarii was used for a South mic characters. The species (until now without American species, although its type locality a type specimen) is the only representative of was Java. the genus in Europe and has been described Corner (1994), whose field experience numerous times (see below). emphasized Southeast Asian locales from Singapore to Kinabalu, spent relatively little Materials and Methods time in South America, but a few collections were gathered in Brazil. When the Brazilian All microscopic observations were made Oudemansiella specimens were compared to using an Olympus BX60 microscope equipped those from Pacific landmasses, he concluded with Phase Contrast (PhC) optics. In descrip-

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Fig. 1. Oudemansiella canarii (from Neotype, DED 6886). Basidiomata. × 0.7. tions below, E = spore length divided by spore Höhnel. 1909. Sitzungber. Kaiserl. Akad. width; Em = median E of a population of Wissenschaft. Wien 118: 276. spores; Lm = median spore length over a Neotypus hic design.: INDONESIA, population of spores. TFB = Tennessee field Java, Mt. Halimun Nat. Park., loop trail from book number, assigned to fresh specimens for Cikaniki, ~1000 m elev., 8.I.1999, coll DE documentation, including photos. Colors within Desjardin, DED 6886 (SFSU, BO). quotation marks are from Ridgway (1912); Basidiomata (Fig. 1) several, collybioid colors cited alphanumerically are from or marasmioid. Pileus (20-)40-65 mm diam, Kornerup and Wanscher (1967). Herbarium plano-convex sometimes with a small, broad acronyms are from Index Herbariorum (http:// and flattened central umbo or central shallow sciweb.NYBG.org/science2/IndexHerbariorum. depression, pellucid-striate, glabrous, glutinous asp) to viscid with scattered small flecks on or embedded in gluten; disc gray-orange (5B5) to Results orange-white to light orange (5A3-4; “pinkish buff”) paler toward margin to pale orange- Agaricus canarii Junghühn. Praemissa in white (5A2), with some pale pink-orange tints floram cryptogamicam Javae Insulae, Fasc. 1: overall (6A3-4) or rarely nearly white; super- 82-83 (1838). ficial floccules usually off-white when young ≡ Amanitopsis canarii (Jungh.) Saccardo. and fresh, becoming sordid grayish by maturity; 1887. Syll. Fung. 5: 27 [listed as questionable] margin usually paler than limb or disc, ≡ Oudemansiella canarii (Jungh.) von translucent, sometimes (especially in younger

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Figs 2, 3. Oudemansiella canarii (from Neotype). 2. Hyphae of superficial pileal patches. 3. Pileipellis elements. Below: clavate hyphal terminals. Above: rod-like superficial, disarticulated cells. Bars = 20 µm. individuals), with occasional, small, white, rod-shaped cells 38-80 × 3-6.5 µm, easily triangular appendicula as remnants of ephemeral disarticulated or crumpled so as to appear partial veil. Lamellae horizontal, broadly disarticulated, generally repent at or near sur- adnate to shallowly adnexed, subdistant to face of glutin, the whole system arising from distant with 2-3 series of lamellulae, convex, interwoven outer pileus tramal hyphae as side broad (-5 mm), thick, white when young, branches, thin-walled, without clamp connec- orange white (5A2) to pink white (6-7A2) in tions; contents of subterminal cells homoge- age. Stipe 30-50(-80) mm long, 2-5(-7) mm neous, hyaline. Pileus tramal hyphae inflated broad at apex, 5-8(-13) mm broad at base, up to 27 µm diam, thin- to thick-walled (wall central to eccentric, curved, tough, fibrous, never more than 1 µm thick), with rare clamp solid; surface covered at first with evanescent, connections, constricted at septa; contents homo- buff to pale pinkish white floccules (not part of geneous, hyaline. Anatomy of scattered flecks partial veil), pale pinkish white (7A2) at first, on pileus surface (Fig. 2): typical slender, darkening at apex to gray red (7B4-5), at base cylindrical pileipellis hyphae (without clamp slightly darker, later longitudinally fibrillose, connections) converge and gather into congest- gradually enlarged downward to a subbulbous ed patches of erumpent hyphal chains in which or bulbous base. terminal cells become increasingly inflated, Habitat: Scattered on fallen tree (of eventually shorter and more broadly fusoid Canarium communis Linn., teste Junghühn). than the slender hyphae from which they arise. Pileipellis (Fig. 3) an ixotrichoderm consis- Terminal cells 12-55 × 12-17 µm, firm- to ting of an erect branched hyphal system with thick-walled (wall never more than 1 µm thick); terminal cells often slightly expanded up to 8 contents homogeneous, hyaline singly, slightly µm diam, appearing as though devoid of more pigmented terminally. “H-“ connections protoplasm or with protoplasm coagulated and branched hyphae numerous. Pleurocystidia against cell wall, surmounted by chains of (Fig. 4) arising deep in subhymenium, 242-280

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Figs 4, 5. Oudemansiella canarii (from Neotype). 4. Pleurocystidia. 5. Basidia and basidiospores. Bars = 20 µm.

× 44-50 µm (at widest point), pedicellate, clavate to broadly clavate, conspicuously clam- broadly proximally inflated with extended, ped, thin-walled; contents homo- to heteroge- subcylindrical neck, obscurely clamped, thick- neous, condensed near wall, hyaline. Stipe walled (wall up to 2.5 µm thick over bulb, up surface generally smooth, of repent, thick- to 1.5 µm thick over extended neck); contents walled (wall up to 1.5 µm thick) hyphae. more or less homogeneous in bulb, with Patches of caulocystidia similar to pileus floc- coagulated protoplasm in neck, subrefringent at cules, scattered, small, ill-defined. Caulocystidia apex; individuals from near the lamellar margin (Fig.7) uni- to few-celled, keg-shaped to clavate, generally shorter, 125-160 × 40-47 µm, long- firm- to thick-walled (wall up to 1.5 µm thick), pedicellate, clamped, otherwise as above. without clamp connections on terminal or Basidia (Fig. 5) 78-90 × 25-30 µm, broadly subterminal septa; contents heterogeneous, of clavate to suburniform from a severely pinched ill-defined protoplasm, hyaline. base, obscurely clamped, refringent (PhC) when Commentary: DED’s field drawings of mature, then losing refringency, thick-walled basidiomata point out stipe floccules as well as (wall up to 1.5 µm thick); contents hetero- pileus floccules. In both cases, these appear not geneous, somewhat coagulated by maturity. to be remnants of a partial or universal veil, but Basidiospores (from hymenium; Fig. 5) 19-25 productions of the outermost pileipellis tissues × 18-23 µm (E = 1.00-1.15; Em = 1.06; Lm = themselves. Corner’s (1934) detailed account 21.8 µm), globose to subglobose, irregular in of the partial veil cannot be seen in mature outline, refringent, thick-walled (wall up to 0.5 material but only in very small primordial. µm thick); contents multigranular, developing Junghühn specifically drew attention to an amorphous guttule by maturity. Lamellar what he interpreted as a veil. Some references edge sterile, not significantly extending in follow: “…velo simplici, universali, fugaci…” KOH, a solid beard of cheilocystidia. Cheilo- “…velo universali floccoso-membranaceo e basi cystidia (Fig. 6) 42-72 × 13-29 µm, pedicellate, stipite orto fugaci…” “…Velum universale, e

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Figs 6, 7. Oudemansiella canarii (from Neotype). 6. Cheilocystidia. 7. Caulocystidia. Bars = 20 µm. basi stipitis incrassato surgenes, totum fungum ment, unlike the annulus of O. mucida and primo obtegens, citissime rumpens, deorsum similar basidiomata. subpersistens, volvamque brevissimam ocreae- Pleurocystidia from near lamellar attach- formem tuberculo stipitis adnatum efficiens, ment to pileus are, on the average, longer and sursum vero (in superficie pilei) fatiscens, stouter than those from nearer to the lamellar squamulis s. membranulis adglutinatis hinc edge. Apices of the latter are often unextended, inde residuis sparsis.” It is likely, however, so the pleurocystidia appear pedicellate and that Junghühn saw only dried material and broadly fusoid rather than ten pin-shaped or interpreted the superficial flecks on pileus and with neck even more extended. perhaps on stipe as evidence of a previous veil. Corner (1934) studied ontogeny of basi- Corner’s (1934, as Collybia apalosarca) diomata in Collybia apalosarca, which he later detailed description of basidiome ontogeny in equated (1994: 49) with Oudemansiella canarii. what he later considered to be O. canarii In the earlier paper, based on specimens from (Corner, 1994) included an account of the veil, Singapore, he was able to furnish figures which while universal at an early stage, is quickly (1934: Figs. 3, 4) of pileipellis structures which evanescent. But even Corner understood the clearly equate to his later description. He scattered flecks on pileus surface to be vestigial interpreted the terminal cells as erect, emerging velar tissue. Similarly, Berkeley and Curtis through a slimy matrix. Our observations are (1868) thought their Agaricus cubensis was from sections and the orientation of the pilei- similar to Amanita gemmata through just such pellis tissue could not be accurately concluded. scattered, raised flecks, and Saccardo concluded The question is whether these chains of cells his incorrect placement of A. canarii in Amani- are erect [as figured by Corner (1994: 64, Fig. topsis probably on Junghühn’s description of 6 for O. lianicola; 66, Fig. 7 for O. platensis] the volva-like veil. Interestingly, however, or whether they are decumbent, as in Megacol- Junghühn did not mention a viscid or glutinous lybia and Clitocybula. Baroni and Ortiz (2002) pileus surface, a feature shared by all basidio- furnished the clearest illustration of this mata examined by us. Corner (1934) was able pileipellis, showing cells of the pileipellis itself to record basidiome ontogeny including presence and of a portion of one of the superficial of a fugacious veil. All evidence for this veil floccules (specimen Baroni 8001, cited below). disappeared very early in basidiome develop- Traditionally, O. canarii has been

86 Fungal Diversity considered pantropical, including New and Old sylvatica (usually standing, occasionally reclin- World tropics as well as Africa. Corner’s ing) that most mycophiles recognize them on (1994) recent contribution questioned this sight. Several recent and classic works include premise, separating O. canarii from O. platensis it under one of its generic names. For example, (South America) and adding some new species Walvogel et al. (2001), Breitenbach and Kränzlin from the Old World (i.e. O. lianicola Corner, (1991), Lange (1935), Kühner and Romagnesi O. crassifolia Corner, O. submucida Corner). (1953), Kreisel (1987) and Boekhout (1999) all In addition to the neotype specimen of O. canarii take up the organism and Fries (1821) wrote (see above), additional basidiomata with “In truncis vetustis, apprime Fagi, vulgaris, identical disarticulated ixotrichoderm, superficial Jul.-Dec. (v.v.)” Fagus was present only in floccules, pleuro- and cheilocystidia have been southern Sweden at the time of his writing of seen from tropical and subtropical New World Systema Mycologicum. locations. Thus, at least as far as morphology can be employed, O. canarii is present in the Agaricus mucidus Schrader. Spicilegium Florae New World. Germanicae, pars prior: 116 (1794). New World specimens of O. canarii examined: ≡ Agaricus mucidus Schrader: Fries. MEXICO, Veracruz, Mpio.de Cajete, Plan de Cedeño, Syst. Mycol. 1: 28 (1821). 5.VI.1981, coll F Ventura (as Oudemansiella), Ventura 18568 (FCME). UNITED STATES, Florida, Alachua ≡ Armillaria mucida (Schrad.: Fr.) P. Co., Gainesville, San Felasco State Preserve, Kummer.: 135 (1871). 6.VIII.1992, coll TJ Baroni, J Kimbrough, S. Both (as O. ≡ Collybia mucida (Schrad.: Fr.) Quélet. canarii), TFB 6865 (CORT); same location, 8.VIII.1996, Enchirid. Fung.: 27 (1886). coll TJ Baroni, TJB 8001 (CORT); same location, ≡ Lepiota mucida (Schrad.: Fr.) Schröter 19.VIII.1996, coll TJ Baroni, TJB 8083 (CORT). Marion Co., Ocala Nat. For., Juniper Springs Nature Trail, in Cohn. Kryptog.-Fl. Schlesien 3(1): 671 7.XII.1998, coll KWH, TFB 10230 (TENN 57140). (1889). Texas, Hardin Co., Saratoga, Lance Rosier Unit, Big ≡ Mucidula mucida (Schrad.: Fr.) Thicket National Preserve, Teel Rd at Cypress Swamp, Patouillard. Hymenomy. Eur.: 96 (1887). 14.VI.2007, coll Eastfield College Class, det DP Lewis, ≡ (Schrad.: Fr.) v. DPJ 7927 (TENN). Höhnel. Sber. Akad. Wiss. Wien 119: 9

[Fragm. Mykol. 12] (1910). Agaricus mucidus Schrader: Fries. Syst. Neotypus hic design.: GERMANY, Mycol. 1: 28 (1821). Thuringia, Unstrut-Hainich Co., vic. Craula, Although the species epithet, mucidus (in Hainich Nat. Park, 51º4′58.84″ N, 10º30′51.89″ Agaricus), has a nomenclatural starting date of E, elev 1135 ft, 16.X.2007, coll S. Kunnert & J. 1821 (Fries, 1821), the name was adopted by Wilhelm, comm. D. Krüger, TENN 62246. Fries from the epithet proposed by Wilhelm Pileus -40 mm broad, shallowly convex Adolph Schrader. Schrader (see Stafleu and with inrolled margin (including appendiculate Cowan, 1985; Daniels and Stafleu, 1974) was tissue), randomly wrinkled (not radially), off- resident and director at the Göttingen, white overall when young, gradually darkening Germany, Botanical Gardens, and although his with age to plumbeo-fuscous, but always with Spicilegium Florae Germanicae was published transparent glutinous surface so as to appear as in 1794, he was a contemporary of Christiaan porcelain, usually darker over disc, somewhat Hendrik Persoon (1801), who also adopted the lighter toward margin, smooth; margin signifi- epithet. In order to honor taxonomic history, cantly appendiculate when young, with edge therefore, it is necessary to seek a type attached to annulus at about half way between specimen for Schrader’s epithet from the lamellar extent and stipe surface; appendiculate environs of Göttingen. margin viscid when fresh, dark cream buff. A collection of A. mucidus was gathered Lamellae adnate with significant decurrent tooth, by state forest personnel in Thuringia, passed subdistant, white to pale buff, without conspi- to DK, thence to RHP, and is designated here cuous pleurocystidial necks; margin even, not to serve as neotype. A detailed description of marginate. Stipe longitudinally lined, silky- the specimen follows only to unmistakably link striate, off-white or cream above annulus when the epithet to a circumscription. Basidiomata of fresh, downward increasingly gray to gray- this are so common in Europe on Fagus

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Figs 8, 9. Oudemansiella mucida (from Neotype). 8. Pileipellis elements. A. From pileus disc. B. From pileus margin. 9. Pleurocystidia. Bars = 20 µm. brown, more or less equal, expanded at base layer thick, of tightly interwoven, strangulate, into an obpyriform or pad-like base on bark; slender (never more than 3 µm diam), hyaline, annulus tightly tomentose, off-white when fresh, thin-walled, frequently clamped hyphae obvious, flairing or pendant, cream-colored on producing basidia of different ages (stages of upper surface, developing deep grey on lower maturity), often as digitate, uninflated terminal surface by maturity, often minutely papillose cells. No inflated tramal cells seen. Basidia on lower surface near margin. (Fig. 10) 65-100 × 19-23 µm, subcylindrical Pileipellis over disc (Fig. 8A) embedded when immature, clavate when mature, distinctly in viscid matrix, constructed of coralloid- clamped (especially evident in young indivi- branched processes, with apices occasionally duals), with slightly pinched base, 4-spored; inflated into small pileocystidia. Hyphae 3-6 contents coscinoidal to multigranular when µm diam, thin-walled, occasionally conspicu- immature, with scattered large guttules by ously clamped (but usually not so), hyaline. maturity, refringent (PhC). Basidiospores (Fig. Pileocystidia 18-32 × 6-8 µm, clavate, thin- 10) 16-20 × 15-19 µm (E = 1.00-1.18; Em = walled, not clamped; contents homogeneous, 1.06; Lm = 18.7 µm), globose to subglobose, hyaline. Pileipellis near margin (Fig. 8B) smooth, uniguttulate when mature, thin-walled embedded in viscid matrix, constructed of (but appearing thick-walled as guttule different- coralloid-branched hyphal systems, with apices iates), hyaline; contents uniguttulate when often inflated into clavate pileocystidia. Hyphae mature, refringent (PhC). Cheilocystidia (Fig. 3-6 µm diam, thin-walled, hyaline, occasionally 11) a solid beard, poorly developed, 25-65 × 7- conspicuously clamped. Pileocystidia 20-38 × 11 µm (at widest point), prominently clamped, 9-11 µm, clavate, thin-walled, not clamped; digitate when immature, eventually broadening contents homogeneous, hyaline. Pleurocystidia somewhat apically; contents scattered-multi- (Fig. 9) widely scattered, almost submerged in guttulate at all ages. Stipe surface above annulus hymenium, 84-103 × 34-41 µm, pedicellate, smooth, a thick repent layer of slender hyphae, utriform, thin-walled, probably clamped, hyaline; 2-3 µm diam, thin-walled, rarely but conspi- contents homogeneous, hyaline. Subhymenial cuously clamped, which appear to be free or

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Figs 10, 11. Oudemansiella mucida (from Neotype). 10. Basidia and basidiospores. 11. Cheilocystidia. Bars = 20 µm.

Figs 12, 13. Oudemansiella species. 12. Oudemansiella mucida (from Neotype). Hyphae from annulus. A. Hyphae from upper surface toward annulus margin. B. Hyphae of fibrillose attachment to stipe. C. Hyphae from gray inrolled annulus margin. 13. Oudemansiella platensis (from Epitype). (1) Pilepellis hyphae (2). Portion of hyphal wall shown as stippling is semi-gelatinized. Bars = 20 µm.

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Figs 14, 15. Oudemansiella platensis. Basidiomata. 14. Epitype, × 0.7. 15. DED 8084, Brazil, ×0.7.

90 Fungal Diversity slightly coherent but not strongly so (liberating pileus margin as appendiculae, although too in squashes). Hyphae becoming somewhat small to be seen without a lens. Very soon after thicker (4-6 µm diam) toward stipe medullary disarticulation, these fibrils are adsorbed and tissue but otherwise as surface hyphae. Stipe become virtually invisible. All this was not surface below annulus identical to above, with described by Corner (1934, 1994), who dealt thick superficial layer of slender, frequently with O. canarii, which does not exhibit an and conspicuously clamped hyphae 2-3 µm annulus in mature basidiomata. diam. Hyphae inward somewhat stouter (-7 µm Hyphae of annulus tissues resemble those diam) with semi-gelatinizing walls and rare of the stipe surface both above and below clamps. Upper surface of annulus (Fig. 12A) annulus (slender, strangulate, frequently medal- fibrillose as attached to stipe surface; hyphae lion clamped, thin-walled, hyaline). The only slender (2-4 µm diam), equal, thin-walled, difference between upper and lower annular frequently clamped, similar to those of grey surfaces is that the upper surface is covered inrolled annulus margin. Hyphae of inner with spores. surface (upper surface of annulus away from Horak (1968) included Mucidula, and stipe; Fig. 12B) constructed of slightly wider examined a specimen of M. mucida (France, hyphae (3-5 µm diam, thin-walled, frequently Carnelle (S.-et-O.), X.1898, leg. N. Patouillard clamped, equal, somewhat strangulate). Tissue (FH). The specimen was cited as “lectotype.” of inrolled gray margin of annulus (Fig. 12C) The type specimen for Mucidula mucida, constructed of tightly interwoven, free, slender nomenclaturally, must be the same as that for hyphae (2-5 µm diam, quite equal except for Agaricus mucidus, for Patouillard did not branching areas), frequently and conspicuously propose a new species but only proposed a clamped, hyaline, thin-walled (just like those of transfer from Agaricus to Mucidula with no stipe surface – see above); gloeoplerous hyphae change in typification of the epithet. common, 4-7 µm diam, aseptate, subrefringent. A morphological variant of O. mucida Commentary: All basidiome surfaces are occurs in temperate Asia, differing in less constructed of complex hyphae. Pileipellis and complex pileipellis tissue, larger spores, more hymenium include subpellis and subhymenial prominent pleurocystidia, well-developed cheilo- layers of coralloid, slender, clamped hyphae, cystidia and paler basidiome color. Before and cheilocystidia are produced by identical distinguishing this variant, Petersen and Halling hyphae with conspicuous clamps. Annulus (1994) reported on a series of pairing experi- tissues and stipe surface are composed of very ments in O. mucida. Normally, dikaryons slender, frequently clamped hyphae. produced in pairings among Asian isolates and Corner (1994: Fig. 10, p 71) figured the among European isolates produced abundant pileipellis of O. mucida (as compared to that of clamp connections. In vitro experiments pairing O. submucida). Clavate pileocystidia were monokaryon isolates of Asian collections versus correctly pictured, but the coralloid hyphal monokaryon isolates of European collections system which produces them was not shown. of O. mucida, however, showed sexual Two pilocystidia were shown as fusiform with compatibility when resultant dikaryons were mammilate tip, but we see no such on this (or observed using a nuclear stain, but produced no other) specimen. clamp connections. Examination of the basidio- The structure and attachment of pileus mata from which the monokaryon isolates were margin and annulus has not been accurately derived confirmed that the Asian basidiomata described (to our knowledge) in this species. were of the morphological variant (which will The annulus clearly does not attach to the be proposed elsewhere). The inference from normal pileus margin, but the pileus margin these experiments is that sexual compatibility extends significantly (with no flesh and no was conserved for nuclear compatibility and lamellar extension) and the suture line between nuclear migration (usually ascribed to the pileus margin and annulus is clearly marked. mating type A gene), but not for formation of As disarticulation occurs, minute, delicate clamp connections (usually ascribed to the fibrils of annular tissue remain attached to the mating type B gene).

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Figs 16, 17. Oudemansiella platensis (from Epitype). 16. Pileipellis hyphae terminal cells. 17. Cells and hyphae of superficial pileal flecks. Bars = 20 µm.

Oudemansia platensis Spegazzini. An. Soc. distant, subventricose, -7 mm deep, white, in Cient. Argen. 10(6): 280 (1880). three ranks. Stipe central, bent, silky, “drab ≡ Oudemansiella platensis (Speg.) Speg. gray” upward, at base abruptly expanded, “hair An. Soc. Cient. Argen. 12: 24. (1881). brown;” evidence of partial veil none. Odor ≡ Psalliota platensis (Speg.) Herter, Est. and taste negligible. On hardwood trunk. Bot. Reg. Uruguaya, III Florula Uruguay. Pl. Pileipellis a lax ixotrichodermium com- Avasculares (Montevideo): 43 (1933). posed of chains of 5-8 elongate-fusiform cells Holotypus (design. Spegazzini, 1880): (Fig. 13) ending in a narrowly fusiform, ARGENTINA, Dist. Federal, Palermo, X.1880, fusiform or subclavate terminal cell (Fig. 16), on dead branches of Erythrina cristagalli L. firm-walled (wall never more than 0.5 µm (LPS!) thick); septa apparently rarely clamped, most Epitypus hic design.: ARGENTINA, distinctly not so; contents homogeneous through- Misiones Prov., Iguazú National Park, Isla San out, becoming olive-tan toward terminus with Martin, ca 500 ft elev., 24.V.2001, coll RHP terminal cells most strongly pigmented. Scat- and E Albertó, TFB 10746 (TENN 58954) tered patches or warts on pileus surface Basidiomata (Fig. 14; Fig. 15 of Brazilian composed of a polycystoderm (Fig. 17), the collection DED 8084) five, armillarioid. Pileus proximal cells of which are relatively narrow 35-62 mm broad, shallowly convex, sometimes and long (45-86 × 8-10 µm) becoming more slightly depressed over stipe, viscid, radially and more inflated until terminal cells are rugulose, with scattered, dark gray, small subglobose to sphaeropedunculate (29-48 × 22- patches or pyramidal warts, “light drab” over 44 µm), thin- to thick-walled (wall never more disc and limb; margin off-white, crenulate to than 0.5 µm thick), with septa appearing double; sublacerate; pileus pellicle somewhat receding clamp connections absent; contents homoge- at margin; pileus flesh over stipe pure white, neous, hyaline. Pileus trama over stipe pure soft. Lamellae adnate with decurrent tooth, white, very light weight when dry (as though

92 Fungal Diversity

Figs 18-20. Oudemansiella platensis (from Epitype). 18. Pleurocystidia. 19. Basidia and basidiospores. 20. Cheilocystidia. Bars = 20 µm. composed of wide, loosely packed hyphae); Pleurocystidia (Fig. 18) scattered, obvious on tramal hyphae 5-20 µm diam, long-celled but lamellar surface at 30× in median area of distinctly clamped, relatively loosely packed, lamella (not near margin and not near attach- hyaline, without evidence of any gelatinous ment to pileus), 135-260 × 31-40 µm, narrowly matrix, thick-walled (wall 1-1.5 µm thick). pedicellate, inflated somewhat proximally

93 (ampulliform) then extended into a long, with complex tissue: outermost layer of tightly broadly cylindrical neck and rounded apex, interwoven, white tomentum perhaps the obscurely clamped, thick-walled (wall up to 2 remnant of a partial veil, covering a woody rind µm thick over proximal swelling, then 1-1.5 µm of 2-3 layers of carbuncle tissue somewhat thick over neck); contents homogeneous, grayish; medullary tissue parenchymatous, hyaline. Lamellar trama constructed of two resembling the tissue of Sambucus pith. hyphal types: 1) 10-20 µm diam, long-celled, Hyphae of tomentum over basal bulb similar to broadly cylindrical, constricted somewhat at superficial hyphae just distal to bulb (two septa, obscurely clamped, thick-walled (wall up sentences above), but much longer-celled, to 2 µm thick); contents homogeneous, hyaline; apparently without clamp connections and and 2) 4.5-8.5 µm diam, equal, thin-walled, commonly secondarily septate (“cloissons de rarely septate (but clamped). Subhymenium retret,” not constricted, very thin curtains tightly compressed but remaining filamentous across filamentous cell), firm- to thick-walled (not parenchymatous), with abundant clamp (wall never more than 1 µm thick). connections. Basidia (Fig. 19) 68-80 × 23-29 Commentary: Corner (1994: 65-66) µm, broadly clavate from a pinched base, (2-)4- reported that his examination of Oudemansiella spored, obscurely clamped, thin- to thick- specimens from three areas of Brazil showed walled (wall never more than 0.7 µm thick); that they differed from the common southeast contents homogeneous when immature to Asian taxon he had collected in Malaysia. His heterogeneous/multiguttulate by maturity; conclusion was that O. platensis differed from sterigmata very stout, curved. Basidiospores O. canarii, and he contrasted some micro- from pileus surface (Fig. 19) 18-22.5 × 17-21 characters. But his conclusions were based on µm (E = 1-1.09(-1.14); Em = 1.07; Lm = 20.6 personal experience, because no type speci- µm), globose to subglobose, often irregular in mens were examined (and for O. canarii, none outline as though a loose episporium, distinctly existed). Baroni and Ortiz (2002) reported that puckered when less than mature, refringent, the type specimen of O. platensis was useless thick-walled (wall never more than 0.7 µm (with which we concur), so until the present thick); contents apparently uniguttulate, but paper, adequate type material could not be guttule irregular in outline. Lamellar margin compared. sterile, a solid beard of cheilocystidia. Cheilo- The only illustration of putative O. cystidia (Fig. 20) 38-74(-139) × 11-26 µm, platensis furnished by Corner (1994: Fig. 7, clavate to broadly clavate, conspicuously left) shows a polycystoderm (his term, pseudo- clamped, thin-walled, occasionally pleurocys- parenchyma-like) as contrasted to an imperfect tidial; contents homogeneous or heterogeneous ixotrichoderm for O. canarii (his Fig. 7, right). and then with scattered refringent guttules, The polycystoderm seems evidently to be a sometimes sizable. Upper stipe surface silky; section through a pileus fleck or patch (see our hyphae strictly longitudinal, tightly packed, Fig. 17) as compared to the pileipellis separate coherent (small bits of adherent substance from such a patch (see our Fig. 16). Thus, in loose in mount). Surface hyphae variable in our opinion, Corner’s figures do not equate to width, 3-14 µm diam, hyaline, firm- to thick- our understanding of these structures. walled (wall up to 1 µm thick), sparsely bran- Unfortunately, Corner (1934) did not ched, conspicuously clamped (but long-celled); describe or figure pleurocystidia or cheilocys- widely scattered hyphal tips (caulocystidia) tidia of C. apalosarca (= O. canarii), so those repent to curved-erect, 5-9 µm diam, narrowly structures cannot be equated with the specimen fusiform. Stipe surface near basal bulb similar described above or in his later publication to above, but with very light thatch of super- (Corner, 1994). Later, however, Corner (1994: ficial hyphae; hyphae generally longitudinal, Fig. 3) figured a single pleurocystidium free, 4-11 µm diam, without clamp connec- (among basidia, for O. canarii) which matches tions, firm-walled, easily disarticulated (but the form seen in most Oudemansiella taxa. some adherent substance covering hyphae very It seems clear that the morphospecies O. lightly and stretching over separating septa); canarii is present in New World tropics and “H-“ connections common. Stipe basal bulb subtropics (see aboce under O. canarii).

94 Fungal Diversity Conversely, basidiomata with pileipellis Grand (as O. canarii), TFB 11198 (TENN 59734); San structure similar to that of O. platensis have José de los Matas, vic. Los Montones, La Placeta, N 19°14.433′, W 70°53.882′, 8.I.2003, coll EA Grand (as been examined from other South American O. canarii), TFB 11703 (TENN59771). ECUADOR, locations. Oudemansiella platensis, at least Napo Prov., San Carlos, INIAP – Napa-Payamino Exp. from initial observations, seems limited to the Station, 0°20′ S, 76°50′ W, no date, coll Catherine New World. At this time, no conclusions can Barbour (as O. platensis), Barbour 1 (E 00218109). be put forward on taxa in Africa in spite of Pegler’s (1977) inclusion of O. canarii. He Discussion considered O. platensis to be a synonym under O. canarii, with many other synonyms accord- The presence of O. canarii in the New ingly. World has been verified, where its distribution The report by Petersen and Halling seems to be parapatric with that of O. platensis. (1993) on the mating system of “O. canarii” Conversely, there are no data which suggest was erroneous. Illustrations and descriptions of that O. platensis occurs in the Old World. those basidiomata match the taxonomic Molecular comparisons of collections from characters of O. platensis. various locations will indicate how similar are Additional specimens of O. platensis examined: New and Old World O. canarii. Accurate ARGENTINA, Misiones Prov., Iguazu National Park, descriptions and illustrations from type 24.V.2001, coll RHP and E Albertó, TFB 10746 (TENN material may aid in accurate identificiation. 58954); same location, Hotel Internacional grounds, S 25°40.785′, W 54°26.724′, 29.V.2001, coll KWH, TFB Perhaps a similar example of such possible 11306 (TENN 59012). BRAZIL, Rio de Janeiro, date distribution involves Lentinula boryana (Berk. and location unknown, coll. Glaziou, Glaziou 9145 & Mont.) Pegler, which extends from Brazil to [K(M) 154851!; holotype of Agaricus radiculosus southern United States and L. raphanipes Cooke]; Manaus, 17.X.1948, coll EJH Corner (as O. (Murrill) Mata & R.H. Petersen (also in the ?platensis), s.n. (E 227801); Mato Grosso, Chavantina, 31.I.1968, coll EJH Corner (as O. platensis), s.n. (E United States as well as Caribbean land- 227802). COLOMBIA, Dpto. Antioquia, Municipio de masses). The former was recognized for many Medellin, campus universitario, Universidad de years, but the latter was revealed through Antioquia, 12.X.2005, coll & det Doris Gallo-G, Gallo- pairing experiments and type specimen studies G 100 (NY). COSTA RICA, Alajuela Prov., Bosque el (Petersen et al., 1998; Mata et al., 2001; Mata Niño, Reserva Forestal de Grecia, N 10°08′38″, W 84°14′62″, 27.VI. 1990, coll RHP, TFB 9649 (TENN and Petersen, 2001). A somewhat contrasting 56610); Cartago Prov., Oresi Dist., Paraiso Co., Tapanti pattern has been termed “the subpruinosus National Park, Panmando Trail, 9°44′57″ N, 83°46′54″ pattern” by Petersen and Hughes (2007) and W. 15.VI.1999, coll JL Mata & R Nuñez, TFB 10071 includes the Caribbean, South and/or Central (TENN 58011); Guanacaste Prov., Santa Rosa, National America and tropical Pacific landmasses. Park, 3.VI.1994, coll TJ Baroni, TJB 7450 (CORT); Puntarenas Prov., vic. Golfito, Pavones, Weston rainforest, N 8°15′ W 83°1′, 4.VII.1992, coll RHP & References KWH (as O. canarii), TFB 4886 (TENN 51078); same location, N 8°15′, W 83°15′, 5.VII.1992, coll KWH (as Baroni, T.J. and Ortiz, B. (2002). New species of O. canarii), TFB 4898 (TENN 51190); vic Santa Eelena, Oudemansiella and Pouzarella (Basidiomycetes: Hotel Heliconia, 400 m before entrance to Sta. Elena ) from Puerto Rico. Mycotaxon 82: Reserve, N 10°20′32″, W 84°47′55″. 1500-1600 m elev., 269-279. 17.III.1999, coll JL Mata (as O. canarii), TFB 10059 Berkeley, M.J. and Curtis, M.A. (1868). Fungi Cubensis (TENN 57626); Coto Brus Co., Hacienda La Amistad, N (Hymenomycetes). J. Linn. Soc., London 10: 282. 8°54.218′, W 82°47.401′, 3.VII.1998, coll RHP (as O. Boekhout, T. (1999). Oudemansiella. In: Flora canarii), TFB 9482 (TENN 56510); same location, N Agaricina Neerlandica., vol. 4. (eds. C. Bas, T. 8°54.218′, W 82°47.401′, 4.VII.1998, coll J Cifuentes & Kuyper, M.E. Noordeloos and E.C. Vellinga) RHP, TFB 9906 (TENN 56534); same location, N Balkema, Amsterdam: 177-178 (with drawings). 8°54.218′, W 82°47.401′, 5.VII.1998, coll RHP (as O. Breitenbach, J. and Kränzlin, F. (1991). Fungi of canarii), TFB 9933 (TENN 56559). CUBA, VII.1857, Switzerland, vol. 3. [see p. 308 with illustrations coll. C. Wright, s.n. (FH; holotype of Agaricus cubensis on p. 309]. Berk. & M.A. Curtis); location unknown, Fungi Corner, E.J.H. (1934). An evolutionary study in agarics: Cubenses Wrightiani No. 9, coll. C. Wright, 13.V, Collybia apalosarca and the veils. Transactions Wright 11 (FH, bar code: 00079977; holotype of of the British Mycological Society 19: 39-88. Agaricus cheimonophyllus Berk. & M.A. Curtis). Corner, E.J.H. (1994). On the agaric genera Hohen- DOMINICAN REPUBLIC, Jarabaroa, Salto de buehelia and Oudemansiella. Part. II: Oudeman- Jimanoa, N 19°03.703′, W 70°51.872′, 6.I.2003, coll EA siella Speg. Garden’s Bull, Singapore 46: 49-75.

95 Daniels, G.S. and Stafleu, F.A. (1977). HI-IAPT Petch, T. (1907). Revisions of Ceylon fungi. Annals of portraits of botanists. Taxon 23: 178. the Royal Botanical Gardens, Peradeniya 4: 21- Farr, M.L. (1973). An annotated list of Spegazzini’s 68. fungus taxa. Biblioth. Mycol. 2: 824-1661. Petch, T. (1910). Annals of the Royal Botanical Gardens, Fries, E.M. (1821). Systema Mycologicum. Vol. 1. Peradeniya 4: 373-439. Gryphswaldiae. Petersen, R.H. and Halling, R.E. (1993). Mating systems Hennings, P. (1900). Hymenomycetinae. In: Die in the Xerulaceae. X. Oudemansiella. Natürlichen Pflanzenfamilien, vol. 1, Abt. 1. (eds. Transactions of the British Mycological Society A. Engler and K. Prantl). Leipzig: 105-313. 34: 409-421. Horak, E. (1968). Synopsis Generum Agaricalium. Petersen, R.H. and Hughes, K.W. (2007). Some agaric Beitrage der Kryptogamenflora de Schweiz 13: 1- distribution patterns involving Pacific landmasses 741. and Pacific Rim. Mycoscience 48: 1-14. Kornerup, A. and Wanscher, J.H. (1967). Methuen Petersen, R.H., Hughes, K.W. and Mata, J.L. (1998). handbook of colour. 2nd edn. Methuen Co., Lentinula boryana intersterility groups and RFLP London. analysis. Inoculum 49: 41. Kreisel, H. (1987). Pilzflora der Deutchen Demokratischen Rick, J. (1905). Pilze aus Rio Grande do Sul. Annales Republik. G. Fischer Verlag, Jena. [see p. 174]. Mycologici 3: 235-240. Kühner, R., and Romagnesi, H. (1953). Flore analytique Ridgway, R. (1912). Color standards and color nomen- des Champignons supérieurs. Masson et Cie, clature. Publ. Privately, Washington, D.C. Paris. [see p. 95, with drawings]. Saccardo, P.A. (1887). Sylloge Fungorum, vol. 5: 1- Lange, J.E. (1935). Flora Agaricina Danica. [reprint 1146. Padua. edition, M. Candusso, 1993] Vol. 1: 400 pp + 104 Singer, R. (1952) [“1950”]. Type studies on pls. [see p. 65 + pl. 41]. Basidiomycetes. Lilloa 23: 147-246. Mata, J.L. and Petersen, R.H. (2001). Type specimen Singer, R. (1964). Oudemansiellinae, Macrocystidiinae, studies in New World Lentinula. Mycotaxon 79: Pseudohiatulinae in South America. Darwiniana 217-229. 13:146-190. Mata, J.L., Petersen, R.H. and Hughes, K.W. (2001). Spegazzini, P.A. (1880). Sobré la Oudemansia platensis. The genus Lentinula in the Americas. Mycologia Annales Societe Cien. Argentina 10: 279-280. 93: 1102-1112. Spegazzini, P.A. (1881). Fungi Argentini additis nonnullis Pegler, D.N. (1977). A preliminary agaric flora of east Brasiliensibus Montevideensibusque. Pugillus Africa. Kew Bulletin Additional Series 6: 1-615. quartus. An. Soc. Cien. Argent. 12: 13-30. Pegler, D.N. (1986). Agaric flora of Sri Lanka. Kew Stafleu, F.A. and Cowan, R.S. (1985). Taxonomic Bulletin, Additional Series 12: 1-519. Literature, 2nd edn., vol. 5: 1-1066. Persoon, C.H. (1801). Synopsis Methodica Fungorum. Waldvogel, F., Neukom, H.P. and Winkler, R. (2001). Göttingen. Pilze Champignons Fungi. AT Press. [see p 374 with photo]

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