28e Jaargang No 1 -4 27 november 1958 FUNGUS · oder Schiffnerula v. Höhn, besitzen bitunicate...

28e Jaargang No 1-4 27 november 1958

FUNGUS O F F I C I E E L O R G A A N V A N D E N E D E R L A N D S E

M Y C O L O G I S C H E V E R E N I G I N G REDACTEUR: Dr R. A. MAAS GEESTERANUS * RIJKSHERBARIUM * LEIDEN

DIE SYSTEMATISCHE STELLUNG DER ASCOMYCETENGATTUNG

ARMATELLA THEISS. ET SYD.

J. A. VON ARX Phytopathologisches Laboratorium “Willie Commelin Scholten", Baarn

(Mit I Abbildung im Text)

Nach der Typusart beurteilt gehört die Gattung Armatella Theiss. et Syd. zu den Meliolaceae und zeichnet sich durch hantelförmige, beidends keulig verdickte, bis zur Reife einzellig bleibende, erst bei der Keimung septierte Ascosporen aus. Die Meliolaceae besitzen Asci mit einer einfachen, zarten Membran und die Perithecien sind mit einem vorgebildeten Mündungsporus versehen. Die Familie gehört zu den Ascohymeniales in die Reihe der Sphaeriales.

Die Ascomycetengattung Armatella wurde von Theissen und Sydow (1915) auf Dimerosporium litseae P. Henn. begründet und innerhalb der Dothideales zu den Polystomellaceae gestellt. Armatella litseae als einzige Art wurde von den genannten Autoren in

mehrfacher Hinsicht falsch charakterisiert. Vor allem sollte der Pilz in der Wirtspflanze ein Hypostroma ausbilden; in Wirklichkeit entstehen die Fruchtkörper auf einem oberflächlichen Mycel. Dies erkannte Hansford (1946), der die Gattung zu den Meliolaceae neben Wageria Stev. et Dalbey [= Balladynopsis Theiss. et Syd. sensu Petrak, 1951] stellte. Kurz charakterisiert wurde Armatella litseae von Hansford und Thirumalachar (1948) nach einer auf Neolitsea zeylanica in Südindien gesammelten Kollektion. In der gleichen Publikation wurde eine auf Cinnamomum zeylanicum gefundene Form als Armatella cinnamomi neu beschrieben. Asci konnten diese Autoren in den beiden Kollektionen nicht beobachten ; über den Bau der Fruchtkörper äusser- ten sie sich nicht.

Von Armatella litseae wurde eine Probe des Originalexemplares untersucht. Bei diesem wächst der Pilz auf Blättern von Litsea glauca und es wurde in Japan gesammelt. Ferner wurden mehrere andere, in Japan und auf den Philippinen gefundene Kollektionen nachgeprüft. Bei den erstgenannten wuchs der Pilz ebenfalls auf Litsea glauca oder auf Cinnamomum pedunculatum, bei den letztgenannten auf Neolitsea-Arten. Das Material stammte grösstenteils aus dem

Naturhistoriska Riksmuseum, Botaniska avdelningen in Stockholm. Alle aus Japan stammenden Kollektionen stimmten gut miteinander überein. Nach

2 FUNGUS [JAARG. 28 No. 1-4

einer von K. Hara auf Litsea glauca gesammelten wurden Abb. 1 und folgende Diagnose entworfen :

Armatella litseae (P. Henn.) Theiss. et Syd.: Querschnitt durch ein reifes Perithecium, gekeimte Ascospore und Mycel mit Hyphopodien.

Die sich epiphyll entwickelnden Mycelrasen sind rundlich oder unregelmässig, 5-10 mm gross, dunkel und

bestehen aus strahlig verlaufenden, reich verzweigten und septierten, braunwandigen, 5-6 μ dicken Hyphen.

Diese sind wechselständig mit braunen, auf einer 5-7 μ grossen Stielzelle sitzenden, im Umrisse rundlichen,

lappig eingebuchteten, 10-15 μ grossen Hyphopodien besetzt. Die sich in den Rasen zerstreut oberflächlich

entwickelnden Perithecien sind niedergedrückt kugelig oder halbkugelig, schwarz, sitzen mit flacher Basis

dem Blatte auf und erreichen einen Durchmesser von 180-280 μ. Ihre Wand ist 27-40 μ dick,

pseudoparenchymatisch und besteht aus 4-6 Lagen von derb- und braunwandigen, aussen rundlichen, innen

etwas flachen, 6-12 μ grossen Zellen. Aussen wittert sie schollig rauh ab. Im Scheitel befindet sich ein

rundlicher, flacher, 18-24 µ weiter, mit Periphysen besetzter Mündungsporus. Die wandständigen, kleineren

hyalinen Zellen aufsitzenden Asci sind oblong oder keulig, gestielt, 60-80 x 22—30 μ gross, besitzen eine

einfache, zarte Membran ohne Apikalstrukturen und enthalten meist 4 Ascosporen. Sie werden von den

fadenförmigen, septierten, hyalinen, 2-3 μ breiten Paraphysen überragt. Die Ascosporen sind länglich,

hantelformig, in der Mitte dünner, beidends keulig verdickt und dann abgerundet. Auch reif ausserhalb der

Asci sind sie hyalin oder nur schwach bräunlich, einzellig, enthalten ein körniges Plasma und messen 35-42 x 11-14 μ. Auf dem Blatte liegende, gekeimte Ascosporen sind braun und in der Mitte septiert, eine der Zellen ist

geschrumpft, die andere hat auf einer Stielzelle ein gelapptes Hyphopodium (Appressorium) gebildet.

Nov. 1958] J. A. von Arx: Armatella Theiss. et Syd. 3

Wie aus dieser Beschreibung und aus Abb. 1 hervorgeht ist Armatella litseae ein eigenartig gebauter, unitunicater Ascomycet, der zu den Ascohymeniales im Sinne von Nannfeldt (1932) gestellt werden muss. Der Pilz steht ziemlich isoliert, ist aber ohne Zweifel am nächsten mit Meliola Fr. verwandt. Wie die Untersuchung verschiedener Arten dieser Gattung zeigte und wie

schon aus den Untersuchungen und Abbildungen von Gaillard (1892), Arnaud (1918) und Graff (1932) hervorgeht, sind die Perithecien bei Meliola ähnlich wie bei Armatella gebaut. Sie besitzen ebenfalls eine scheitelständige, vorgebildete, mit Periphysen besetzte Mündung und die Membran der Asci ist einfach, zart und verschleimt leicht. In ihnen werden ebenfalls vier Sporen angelegt, von denen aber nur zwei zur Reife gelangen, während die beiden anderen frühzeitig degenerieren.

Die Familie der Meliolaceae gehört zu den Ascohymeniales in die Reihe der Sphaeriales, nimmt aber dort eine ziemlich isolierte Stellung ein. In die Familie gehören ausser Armatella die Gattung Meliola und ihre nächsten Verwandten wie Irene Theiss. et Syd. [= Asteridiella McAlpine, vgl. Hansford, 1956] und Amazonia Theiss., alle mit bei der Reife zweisporigen Asci und 4-5 zelligen, braunen Ascosporen. Alle anderen, z.B. von Hansford (1946) zu den Meliolaceae gestellten Gattungen wie Balladyna Rac. oder Schiffnerula v. Höhn, besitzen bitunicate Asci und müssen zu den Ascoloculares und zwar in die Reihe der Dothiorales in die Nähe der Asterinaceae gestellt werden.

ZITIERTE LITERATUR

ARNAUD, G. 1918. Les Astérinées I (Thèse, Montpellier, Coulet et Fils). GAILLARD, A. 1892. Le genre Meliola (Thèse, Paris, Klincksieck). GRAFF, P. W. 1932. The morphological and cytological study of Meliola circinans. In Bull. Torr. bot. Club 59: 241-

266. HANSFORD, G. G. 1946. The foliicolous Ascomycetes, their Parasites and Associated Fungi. In Mycological

papers, G. M. I. Kew, No. 15. ----------- 1956. Tropical Fungi VI. In Sydowia 10: 41-100. ----------- und M. J. THIRUMALACHAR, 1948. Fungi of South India. In Farlowia 3: 285-314. NANNFELDT, J. A. 1932. Studien über die Morphologie und Systematik der nicht-lichenisierten, inoperculaten

Discomyceten. In Nova Acta reg. Soc. Sci. upsal. ser. 4, 8: 2. PETRAK, F. 1951. Ueber die Gattungen Balladynopsis, Wageria und Balladynastrum. In Sydowia 5: 426-428. THEISSEN, F. und H. SYDOW, 1951. Die Dothideales. In Ann. mycol. 13: 149-746.

UEBER DEN ASCOMYCETEN RHAGADOLOBIUM CUCURBITARUM

(REHM) THEISS. ET SYD.

J. A. von Arx

Phytopathologisches Laboratorium “Willie Commelin Scholten”, Baarn

(Mit 2 Abbildungen im Text)

Rhagadolobium cucurbitarum (Rehm) Theiss. et Syd. bildet auf der Unterseite lebender Blätter von Cucurbitaceae ein oberflächliches, farbloses Mycel mit Hyphopodien. Ueber den Hyphen entstehen krustenförmige Stromata; deren radiäre Deckschicht reisst bei der Ascusreife über den Loculi unregelmässig auf. Der Pilz ist eine stromatische Asterinaceae und wird in eine neue Gattung Asterotexis gestellt.

Anlässlich einer Nachprüfung der von Theissen und Sydow (1915) zu den Parmulariaceae und den Polystomellaceae gestellten Pilze wurde auch Rhagadolobium cucurbitarum (Rehm) Theiss. et Syd. untersucht. Dieser ursprünglich als Dothidella cucurbitarum Rehm beschriebene Pilz wurde von den genannten Autoren als zweite Art in die Gattung Rhagadolobium P. Henn. et Lind, gestellt. Rh. hemiteliae als Typusart der Gattung ist ein Parasit auf Farnwedeln und wurde auf den Samoa-Inseln gesammelt. Der Pilz besitzt ein interzelluläres, braunes, hyphiges oder parenchymatisches Hypostroma. Von diesem aus brechen zahlreiche Hyphenbündel durch

die Spaltöffnungen hervor und bilden oberflächlich ein krustenförmiges Stroma mit einer radiär gebauten Deckschicht, unter der sich die Asci und Ascosporen in unregelmässigen Loculi entwickeln. Dieser Pilz ist eine typische Parmulariaceae.

Von Rh. cucurbitarum wurden eine Probe des Originalexemplares und zwei von Sydow in Südamerika gesammelte, in den Sammlungen des Institutes für spezielle Botanik der Eidgen. Technischen Hochschule in Zürich bewahrte Kollektionen untersucht. Bei diesen wuchs der Pilz auf Cucurbita pepo L. ; nach den Angaben von Ciferri (1957) kommt er aber auch auf anderen Cucurbitaceae vor und scheint in Zentral- und Südamerika weit verbreitet zu sein.

Rh. cucurbitarum weicht von Rh. hemiteliae in mehrfacher Hinsicht ab und muss in eine eigene Gattung gestellt werden. Der Pilz ist überhaupt keine Parmulariaceae. Er besitzt ein oberflächliches, farbloses Mycel mit Hyphopodien, die von Theissen und Sydow (1914) als Konidien beschrieben wurden, während sie das Mycel übersahen. Die Stromata entwickeln sich völlig oberflächlich. Die Angabe der genannten Autoren, nach der die Stromata in ihrer ganzen Breite der Epidermis eingewachsen und durch interzellulär ins Blatt eindringende Hyphenzüge

verankert sein sollen, beruht auf einem Beobachtungsfehler. Nach den untersuchten Exemplaren wurde folgende Diagnose entworfen (vgl. auch Abb. 1

und 2).

Die Stromata entwickeln sich regelmässig und ziemlich dicht zerstreut auf der Unterseite der Blätter

zwischen den Borstenhaaren des Wirtes. Sie sind als 2-4 mm grosse, dunkle, unscharf begrenzte Krusten

wahrnehmbar. Das oberflächliche Mycel besteht aus verzweigten und septierten, 4-5 μ breiten, farblosen

Hyphen. Die wechselständigen Hyphopodien sind einzellig, ellipsoidisch oder länglich, braun, 7-1ο μ lang und

5-7 µ breit. Seitlich am Mycel entstehen ferner zahlreiche, rundliche, nur schwach oliven-

Nov. 1958] J. A. von Arx: Rhagadolobium cucurbitarum 5

bräunlich gefärbte, vielzellige Bulbillen, die sich über dem Mycel ausbreiten und seitlich miteinander

verschmelzen. Die so entstehenden, 0.5-4 mm grossen Stromata sind im Umrisse sehr unregelmässig und

erreichen reif eine Höhe von 60-95 µ. Ihre Deckschicht ist 3-5 μ dick, hell oliven- oder gelbbraun und besteht aus

einer lage von 3-6 μ langen und 2-3.5 μ breiten Hyphengliedern. Deren Wände sind nur nach aussen verdickt und

hellbraun gefärbt. Die 10-14 µ dicke Basalschicht ist hyalin, undeutlich kleinzellig oder hyphig. Im Stroma

befinden sich meist mehrere, im Umrisse rundliche oder unregelmässige, oft seitlich miteinander

verschmelzende Loculi. Ueber ihnen wird die Deckschicht aufgesprengt und lappig zurückgeschlagen oder

abgeworfen. Die dicht parallel nebeneinander stehenden Asci sind bitunicat, dickwandig, 30-44 μ lang und 11-14

μ breit. Sie sind von einer aufquellenden, etwas zähen Schleimmasse bedeckt und enthalten acht fast im obern

Drittel septierte, hyaline, bei der Reife hie und da sehr schwach bräunliche, 12-16 x 4-5 μ grosse Ascosporen.

Abb. 1-2. Asterotexis cucurbitarum (Rehm) v. Arx: 1—Mycel mit Hyphopodien und Bulbillen; 2—Schnitt durch den Rand eines Stromas.

Dieser Pilz ist eine typische Asterinaceae mit sehr dicht entstehenden, bald verwachsenden,

stromatische Krusten bildenden Fruchtkörpern. Von den bestehenden Gattungen der Asterinaceae unterscheidet er sich schon durch die farblosen Hyphen. Ferner wachsen die Fruchtkörper nicht invers unter den Hyphen heran, sondern sitzen diesen auf und die Hypopodien sind unter ihnen bleibend mit sehr kleinen Haustorien im Substrat verankert. Am

nächsten ver-

6 FUNGUS [Jaarg. 28 No. 1-4

wandt ist dieser Pilz noch mit den Vertretern der Gattung Symphaster Theiss. et Syd. (Syn.: Xenostomella Syd.). Nach den Originalexemplaren beurteilt sind hier die oberflächlichen Hyphen aber derb und dunkel gefärbt und die dunklen Stromata entwickeln sich unter diesen und besitzen eine dicke, mehrzellschichtige Deckschicht. Ferner sind die Ascosporen

verhältnismässig gross und färben sich bei der Reife braun. Rhagadolobium cucurbitarum muss daher in eine eigene Gattung gestellt werden, die

folgendermassen charakterisiert werden kann :

Asterotexis v. Arx, nov. gen.

Mycelium liberum praesens, ex hyphis ramosis, septatis, hyalinis, hyphopodiatis brunneis constans; Stromata superficialia, dimidiato-scutata vel crustacea, irregularia, plurilocularia; strato basali indistincte et minute celluloso, hyalino ; strato tegente pellucide olivaceo-brunneo, radiatim contexto, in maturitate stellatim vel rima irregulare dehiscente; asci clavati vel ovati, firme bitunicati, octospori, in massa mucosa inclusi; ascosporae oblongo clavatae, 1-septatae, hyalinae, minutulae.

Mycelrasen klein, unscheinbar, oberflächlich; Hyphen verzweigt, septiert, hyalin oder hell gefärbt, mit

lateralen, einzelligen, braunen Hyphopodien und mit bulbillenartigen, hellen, sich flach ausbreitenden

Körperchen besetzt. Stromata über den Hyphen, durch seitliches Verwachsen dieser Körperchen entstehend,

flach krustenförmig, im Umrisse buchtig oder unregelmässig; Deckschicht radiär, dünn, hell, bräunlich, sich

über den unregelmässig rundlichen Loculi spaltig oder rissig öffnend; Basalschicht undeutlich kleinzellig; Asci

bitunicat, keulig oval, parallel stehend, ohne deutliche Paraphysen, einer Schleimmasse eingebettet, 8-sporig;

Ascosporen keulig-oblong, in der Mitte oder fast im obern Drittel septiert, hyalin oder reif schwach bräunlich,

ziemlich klein.

Als Typusart dieser Gattung hat Asterotexis cucurbitarum (Rehm) v. Arx, comb. nov. [Dothidelia cucurbitarum Rehm in Hedwigia 36: 376.1897 (basionym). — Rhagadolobium cucurbitarum (Rehm) Theiss. et Syd. in Ann. mycol. 12: 275. 1914] zu gelten.

ZITIERTE LITERATUR

CIFERRI, R. 1957. Micoflora Domingensis Exsiccata. In Sydowia 10: 130-180. 1956. THEISSEN, F. und H. SYDOW, 1914. Dothideazeen-Studien II. In Ann. mycol 12: 268-281. 1915. ----------------------------- Die Dothideales. In l.c. 13: 149-746.

THE GENERIC NAMES PROPOSED FOR HYMENOMYCETES—IX *

“Meruliaceae” and Cantharellus s. str.

M. A. Donk

Rijksherbarium, Leiden

This ninth contribution deals with the Meruliaceae in a wide sense and with the generic names based on Cantharellus cibarius Fr.

INTRODUCTION.—This paper forms the ninth part of a series planned to give an annotated nomenclatorial enumeration of all generic names proposed for Hymenomycetes. For some general remarks the reader is referred to the “Introduction to the series” in Part I (in Reinwardtia i: 199-203. 1951) and to a paper entitled “The typification of revalidated names”

(Donk in Taxon 6: 245-256. 1957) in which also the use of ‘per’ instead of ‘ex’ in connection with revalidated names is explained.

DEFINITION.—The name Meruliaceae is here taken in a conservative, inclusive sense. The circumscription is the same as of Merulius Fr. sensu lato, for instance of Killermann (in Nat. Pfl Fam., 2. Aufl., 6: 171. 1928). The group is by no means a taxonomic one: currently most of its species are distributed over the Corticiaceae and the Coniophoraceae emend. Donk. It was not well possible to deal with Merulius Fr. without adding a treatment of the generic names based on Cantharellus cibarius Fr.

ACKNOWLEDGMENT.—I am very much indebted to Mr D. A. Reid, The Herbarium, Royal Botanic Gardens, Kew, for linguistic improvements of the manuscript.

ALPHABETICAL ENUMERATION

Agarico-merulius Haller, Enum. meth. Stirp. Helv. indig. 1: 33. 1742 (pre-Linnean name). — This genus was introduced for a species I would be inclined to identify with Merulius tremellosus Schrad. per Fr. —Typonyms: Merulius Fr. (1821; not Merulius [Haller] St.-Am.) and

Trabecularia Bonord. (1857).

Alectorolophoides Earle in Bull. N. York bot. Gdn 5: 407. 1909. — ETYMOLOGY: αλέκτωρ, —

opος, cock; λόφος comb; —ειδής, like. Gender: f. — TYPE SPECIES (by original designation) :

Cantharellus cibarius Fr. — PROTONYM : "Alectorolophoides" ; Batt., Fungi Agri arimin. Hist. 39.

1755.—The names of three species of Battarra’s class IX begin with this word ; all represent

Cantharellus cibarius. I assume that Earle regarded as type the first species, Alectorolophoides

scrobiculatum Batt. (pl. 14 f. A), the first identifiable species with a

* Part I (“Cyphellaceae”) of this series was published in Reinwardtia 1: 199-220. 1951; Part II (Hymenolichenes), in Reinwardtia 2: 435-440. 1954; Part III (“Clavariaceae”), in Reinwardtia 2: 441-493. 1954; Part IV (Boletaceae), in Reinwardtia 3: 275-313. 1955; Part V (“Hydnaceae”), in Taxon 5: 69-80, 95-115. 1956; Part VI (Brachybasidiaceae, Cryptobasidiaceae, Exobasidiaceae), in Reinwardtia 4: 113-118. 1956; Part VII (“Thelephoraceae”), in Taxon 6: 17-28,68-85, 106-123. 1957; Part VIII (Auriculariaceae, Septobasidiaceae, Tremellaceae, Dacrymycetaceae), in Taxon 7 (in press).


‘binomial’ ( = biverbal) name. One is reminded of the fact that the first words of Battarra’s specific names are not comparable to the generic names of De Tournefort’s or Linnaeus’s systems of nomenclature. — SCOPE. ‘‘These are the typical chanterelles. It is unfortunate that the familiar generic name of Cantharellus is antedated.”—Earle (l.c.). Such genera as Dictyolus Quél. (see

“Cyphellaceae”), Merismodes Earle (see “Cyphellaceae”), and Turbinellus Earle were excluded from Cantharellus Adans. per Fr. when the name Alectorolophoides was validly published for the residue. — REMARK. The name was ascribed to Battarra but Earle’s use of it amounts to the introduction of a new generic name, based on the type species he designated, rather than on Battarra’s corresponding species. In this case the two are conspecific. — TYPONYMS: Cantharellus

Adans. per Fr. (Jan. 1, 1821) and Merulius [Haller] St.-Am. (Apr. 1821; preoccupied).

Cantarellus.—See Cantharellus.

Cantharellus Adans. (“Chanterel”) per Fr., Syst. mycol. 1: 316. 1821. — ETYMOLOGY:

Chanterelle, a French vernacular fungus name. Gender: m. — TYPE SPECIES (selected): Fungus sp. Vaill. pl. 11 fs. 14, 75 = Agaricus chantarellus L. = Cantharellus cibarius Fr.—DEVALIDATED NAME:

Chanterel Adans., Fam. PI. 2: 11. 1763.—Adanson’s generic description runs: “Chapeau turbiné concave en-dessus, doublé en-dessous de nervures ou lames rameuses. Porté sur une tige centrale.” He cited specifically ‘‘Fungus. Vaill. Bot. t. 11. f. 9 à 15. Pocillaria. Brown, t. 15. f. 1.” Vaillant’s species are now determined as Cantharellus infundibuliformis (Scop.) per Fr., not Elvella tubaeformis Schaeff., (Vaillant’s figures 9, 10; cf. Fries, Hym. europ. 458. 1874), Craterellus sinuosus (Fr.) Fr. (Vaillant’s figures 11-13; cf. Fries, Hym. europ. 631. 1874), and

Cantharellus cibarius (Vaillant’s figures 14, 15). The inclusion of Pocillaria P. Browne, q.v., is apparently due to a misinterpretation because its gills do not answer to Adanson’s generic description. — The first to use the modern form of the name was De Jussieu: Cantharellus Juss., Gen. PL 4. 1789 (no author’s citation). No species were mentioned, but there can be no doubt about the scope of his group. There is little evidence that there is a direct relation between Cantharellus Juss. and Chanterel Adans. However, De Jussieu was well acquainted with Adanson’s work. There is just the possibility that he chose the specific epithet of the name Agaricus chantarellus L. to serve as a generic name (and in this case spelled it differently, and, perhaps, erroneously ‘Cantharellus’). — It was De Jussieu’s name (rather than Adanson’s) that became current afterwards, because it was taken up by Persoon (in Neues Mag. Bot. 1: 106. 1794 = Tent. 26. 1797, as Cantarellus). He indicated De Jussieu as its author and gave the genus about the circumscription that was the one subsequently accepted by Fries (1821). Of his species, Cantharellus edulis Pers. (= A. chantarellus) and A. aurantiacus Wulf, [as C. “aurantius” (Wulf.) Pers.] may be mentioned. In later work Persoon preferred the name Merulius [Haller] Boehmer,

q.v., to Cantharellus. — Of historic importance is Link’s use of the name (in Mag. Ges. naturf. Fr. Berlin 3: 38. 1809). —VALID RE-PUBLICATION. Fries cited Adanson as the author of the name but used De Jussieu’s form Cantharellus ; and further mentioned in connexion with it Persoon (“disp. p. 26”) and Link; the latter

Nov. 1958] M. A. Donk: Generic names ("Meruliaceae”) 9

with a “!” Thus the chain: Fries 1821—Link 1809—Persoon 1794—De Jussieu 1789—Adanson 1763 is established, but it may be doubted whether the last link is firmly connected with the rest, and whether it would not be better to cite the name as ‘Cantharellus Juss. per Fr.,’ rather than as 'Cantharellus Adans. ("Chanterel”) per Fr.’ — SCOPE. Fries’s genus is essentially that of Persoon

(1794), with some additional elements. —TYPIFICATION. The evident choice is the species indicated above. Nearly all authors who indicated a type mentioned this fungus.1 — REMARKS. S. F. Gray (Nat. Arrang. Brit. PI. 1: 636.1821) validly re-published Cantharellus independently of Fries. He did not cite an author, but we may safely accept that he took the name from Persoon [Syn. Fung. 488. 1801; as Merulius sect. Cantharellus (Juss.) Pers.]. He included Merulius cantharellus (L.) Pers. {Cantharellus cibarius) under the name of Cantharellus vulgaris S. F. Gray. — Quélet (FI. mycol. France 38. 1888) called an emended genus, exclusive of Cantharellus cibarius, "Cantharellus, (J. Bauh.), Quel.”1 — VARIANT SPELLINGS: "Chanterel” ; Adans., l.c.—The ‘original’ spelling! Deliberately reintroduced by Murrill (in N. Amer. Fl. 9: 167. 1910), who treated it as a mere variant of the current name.2 In case one would prefer to dissociate Chanterel Adans. from Cantharellus ‘Adans.’ per Fr., Chanterel Adans. per Murrill would become an orthographically different homonym of Fries’s name, and as a homotypic homonym would again acquire the status of a mere variant spelling. — "Cantarellus”; Pers., l.c., 1794 (“Juss.”) ; Wallr., FI. crypt. Germ. 2: xxxviii, 624. 1833 ("Adans.”). — "Chantarellus. Juss.”; Dubois, Méth. épr. Orléans 153. 1803. — "Cantherellus”; Kauffm. in Pap. Michigan Acad. Sci. 5:

124-125. 1926; Agar. Michigan 1: 32. 1928.—Consistently and apparently deliberately used. — "Chanterelle”; Pfeiffer, Nomencl. bot. 1: 580. 1873.—1 Pfeiffer ascribed this spelling erroneously to Leman (in Diet. Sci. nat. 8: 140. 1817) : the latter author correctly wrote Cantharellus (which he placed as a synonym under Merulius), but also gave the French vernacular form "Chanterelle.” — It could be defended that De Jussieu’s form Cantharellus was an unintentional misspelling of the epithet ‘chantarellus’ (cf. Ag. Chantarellus L., Chanterel Adans., and the vernacular name Chanterelle!). —TYPONYMS: Merulius [Haller] St.-Am. (Apr. 1821; non Fr.) and Alectorolophoides Earle (1909).

Cantherellus.—See Cantharellus. Chantarellus.—See Cantharellus. Chanterel.—See Cantharellus.

Chanterelle.—See Cantharellus.

Gyrophana Pat., Cat. rais. Pl cell. Tunis. 53. 1897. — ETYMOLOGY: γυρός, round; φαίνω, I

show. Gender: f. — TYPE SPECIES (selected for basinym): Merulius lacrimans (Wulf.) per Fr. —

BASINYM: Gyrophora Pat. (1887), q.v. — REMARK. A name change of Gyrophora Pat., the latter

name being preoccupied. — TYPONYMS: Serpula (Pers.) per S. F. Gray (1821), Xylomyzon Pers.

(1825), and Xylophagus Link per Murrill (1903).

1 Quélet (FI. mycol. France 37. 1888) was the only dissendent when he excluded C. cibarius to transfer it with some other species to Craterellus Pers. and to retain the name Cantharellus, for, among others, Cantharellus aurantiacus and for such species as are often placed today in Clitocybe (Fr.) Staude.

2 With “Agaricus Chantarellus L.” as type species.


Gyrophora Pat., Hym. Eur. 143. 1887. — ETYMOLOGY: γύρος, round; φέρω, I bear. Gender: f. — TYPE SPECIES (selected): Merulius lacrimans (Wulf.) per Fr.—SCOPE: “G. lacrymans, G. umbrina, etc.”—Patouillard (l.c.). —TYPIFICATION. The one economically important of the two examples mentioned was indicated as type species by Murrill (in J. Mycol. 9: 99. 1903), and adopted by W.

B. Cooke (Gen. Homobas. 42. 1953) ; for the present name as well as for its isonym Gyrophana. — HOMONYM: Gyrophora Ach. (1803; Gyrophoraceae, Lichenes). — ISONYM: Gyrophana Pat. (1897), q.v. — TYPONYMS: Serpula (Pers.) per S. F. Gray (1821), Xylomyzon Pers. (1825), and Xylophagus Link per Murrill (1903). —STATUS. Impriorable on account of the earlier homonym and, therefore, replaced by Gyrophana Pat.

Leucogyrophana Pouzar in Ceskâ Mykol. 12: 32. 1958. — ETYMOLOGY: λευκός, white; the genus

Gyrophana. — TYPE SPECIES (by original designation and only original species) : Merulius

molluscus Fr.

Merulius Fr., Syst. mycol. 1: 326. Jan. 1, 1821. —ETYMOLOGY: “Nomine ad Morchellas (meras- tute cibarias) denotandas veteres usi sunt, huc transtulit Haller.”—Fries (Syst. mycol. 1: 327. 1821). Gender: m. — TYPE SPECIES (selected): Merulius tremellosus Schrad. per Fr. — HISTORICAL

(before 1821). The first publication of the name Merulius to be taken into account in this connexion seems to have been effected by Boerhaave (Index alter PL Hort. Acad. Lugd.-Bat. 1: 13. 1720) ; he included five species referable to the genus now called Morchella [Dill.] Fr. Linnaeus (Syst. nat. 1735) accepted “Merulius B[oerhaave]” (to drop it again in later work), citing “Morchella D[illenius]” (Cat. Pl. ca Gissam nasc. 187. 1719) as a synonym. In this sense the name has never been taken up again. — Merulius Haller, Enum. meth. Stirp. Helv. indig. 1: 33. 1742· — Von Haller made it clear that he founded a new genus for which he took up the pre-existing name Merulius (and of which he indicated the origin). His species are identifiable with Cantharellus cibarius Fr. and C. (Gomphus) clavatus Pers. per Fr. Another genus, Agarico-merulius Haller (op. cit. p. 33), presumably covers what is now called Merulius tremellosus. — Merulius [Haller] Boehmer, Ludwig Defin. Gen. PL 492. 1760; Haller, Hist. Stirp. ind. Helv. inch. 2: 150. 1768; Scop., FI. carniol., Ed. 2, 2: 461. 1772; Pers., Syn. Fung. 488. 1801.—This is the perpetuation of Von Haller’s genus (1742) after the introduction of Linnaeus’s binomial system of nomenclature. It became gradually widened by the inclusion of sessile and even resupinate fungi, emendations for which Von Haller himself (1768) set the example by merging his Agarico-merulius into it. After the adoption of the name by Persoon (1801), Merulius became currently used in this new sense. — Merulius Pers. in Neues Mag. Bot. 1: 106. 1794 (= Tent. 26. 1797).1—For a short period Persoon had his own genus Merulius. This was when he took up Cantharellus Juss. (= Can-tharellus Adans. per Fr.) for Cantharellus edulis Pers. (= C. cibarius), Agaricus

1 And compare De Jussieu (Gen. Pl. 4. 1789): “Pileus parasiticus suberosis (quandoque carnosus terrestris) sessilis aut raro stipitatus, supra laevis, subtus lamellatus, lamellis radiatis. Huc referatur Agaricus alneus L.”

‘Merulius Juss.’ of Dubois (Méth. épr. Orléans 179. 1803) is this genus, which covered the agarics with lateral cap. Persoon’s genus Merulius of 1794 may be an emendation of it.

Nov. 1958] M. A. Donk: Generic names (“Meruliaceae”) 11

aurantiacus Wulf., Cantharellus umbonatus Pers. (Agaricus muscoides Wulf.) and Cantharellus cinereus Pers. The name Merulius thus had become ‘free’ once more and he gave it (without citing an author) to “M. quercinus” = Daedalea quercina (L.) Pers., “M. sepiarius Schrank” = Lenzites (Gloeophyllum) sepiaria (Wulf.) per Fr., and M. umbrinus Pers. (non Fr.). Thus, Merulius Pers.

1794 is the precursor of Daedalea Pers. (see under Daedalea Pers. per Fr., “Polyporaceae”). As already indicated above Persoon soon favoured the more faithful application of Merulius Haller. — In 1801 Merulius [Haller] was divided by Persoon (op. cit.) into three sections (i) 'Cantharellus', pileo integro, (ii) ‘‘Pileo dimidiato submembranaceo” [with Cantharellus muscigenus (Bull.) per Fr., M. tremellosus Schrad., etc.], (iii) ‘Serpula’ [with M. destruens Pers. = M. lacrimans (Wulf.) per Fr., &c.], and (iv) ‘Gomphus’ (with M. clavatus Pers.). One sees that here is the basis for several later genera: for instance section (i), (ii) in part (exclusive of M. tremellosus), and (iv) taken together equal Fries’s genus Cantharellus Adans. per Fr. ; and M. tremellosus and section (iii) equal the genus Merulius Fr. ; section (ii) in part (exclusive of M. tremellosus) became the later genera Leptoglossum P. Karst, and Leptotus P. Karst., which together equal Corniola S. F. Gray, Dictyolus Quél. ; and section (iv) became Gomphus Pers. (Gomphora Fr.). — SCOPE. The first valid re-publication of Merulius occurred in the starting-point book. Fries’s genus compares with Merulius sect. Serpula Pers. plus such species as M. tremellosus (see above). He listed ten species, of which M. tremellosus is the first. — TYPIFICATION. The type species of Merulius Haller, Cantharellus cibarius, cannot be considered

for Fries’s generic name, because he excluded it. The species unanimously selected today is Merulius tremellosus Clements & Shear (Gen. Fungi 347. 1931), Bondarzev & Singer (in Ann. mycol. 39: 48.1941 ; apud Sing, in Mycologia 36:67.1944), Cunningham (in Bull. Pl. Dis. Div., Dept. sci. ind. Res., N. Zeal. No. 83: 2. 1950), Bondarzew (Trutov. Griby 51. 1953), W. B. Cooke (Gen. Homobas. 62. 1953). It was the only (Finnish) species retained in the genus by Karsten (in Bidr. Känn. Finl. Nat. Folk 48: 341. 1889). The other species of common occurrence and of economic importance that deserves consideration in this regard as a possible rival of M. tremellosus is M. lacrimans (Wulf.) per Fr. It has been selected as type species of generic names introduced for about the same genus as Merulius Fr. : Xylophagus Link, Serpula (Pers.) per S. F. Gray, Xylomyzon Pers. ; it was excluded when Gyrophana Pat. = Gyrophora Pat. was introduced for it and its closest relatives. — REMARK. Since Von Haller’s and Fries’s name are to be typified differently, the latter should not be burdened with ‘Haller’ in the author’s citation.—MONADELPHOUS HOMONYMS : (i) Merulius [Haller] St.-Am., Fl. agen. 556. Apr. 1821. —Type species (selected): Agaricus chantarellus L. (“cantharellus”) = Cantharellus cibarius Fr.—See page 12. —

(ii) Merulius S. F. Gray, Nat. Arrang. Brit. PI. 1: 636. Nov. (?) 1821 (no author’s citation).— Type species (only species) : Agaricus aurantiacus Wulf. = Merulius aurantiacus (Wulf.) Pers.—See page 12. —TYPONYM: Trabecularia Bonord. (1857).

Merulius [Haller] St.-Am., Fl agen. 556. Apr. 1821. — ETYMOLOGY: see under Merulius Fr. Gender: m. — TYPE SPECIES (selected): Agaricus “cantharellus” L. (chantarellus) = Cantharellus cibarius Fr. — PROTONYM: Merulius Haller, Enum. meth. Stirp. Helv. indig. 1: 33. 1742, not Boerhaave 1720. —


DEVALIDATED MAME : Merulius [Haller] Boehmer, Ludwig Def. Gen. PI. 492. 1760; Pers., Syn. Fung. 488. 1801, not Persoon 1794.—For some historical details see under Merulius Fr. — SCOPE.

AS re-published by Saint-Amans (no author’s citation given) the name covered the same group that De Candolle (Fl. franç. 2: 128. 1805) called “Merulius. Hall. Pers.,” and which in turn is an

only slightly emended version of Merulius Haller as defined by Persoon in his “Synopsis.” The first of Saint-Amans species is Agaricus chantarellus L. = Merulius “cantharellus” (L.) Pers. and one of the others, Merulius lacrymans Wulf. — TYPIFICATION. In view of the history of the generic name as discussed under Merulius Fr., the most acceptable choice is Agaricus chantarellus L. and Earle (in Bull. New York bot. Gdn 5: 390, 407. 1909) already listed that species (as Cantharellus cibarius Fr.) as type of “Merulius Haller ... 1768.” It is here accordingly selected for Merulius [Haller] St.-Am. — REMARKS. Fries’s misapplication (1821) of Merulius Haller, now sanctioned by the Code, was not known to Saint-Amans, and his re-publication of the name is part of the overflow of the pre-Friesian era, like the treatments by Hooker (Fl. scot. 2: 25. May 1821, “Withering”), Mérat (Nouv. Fl. Env. Paris, 2e Ed., 1: 49. June 1821), Purton (App. Midi. Fl. 540. After Sept. 4, 1821, who like Hooker followed Withering’s emendation), and Leman (in Diet. Sci. nat. 30: 174-179. 1824). Compare also Roques (Phytogr. médic. 53. Aug. 1821 ; no accompanying generic description). — Persoon (Mycol. europ. 2: 11. 1825) heavily restricted the genus by excluding Gomphus Pers. [per S. F. Gray] and Xylomyzon Pers. (with the scope of Merulius Fr.). The first appearance of this narrow Persoonian concept is to be found in Persoon’s

“Traité sur les Champignons comestibles” (pp. 43, 228. 1818, but not on p. 99). — Many years afterwards O. Kuntze (Rev. Gen. PI. 2: 861. 1891) deliberately again substituted Cantharellus Adans. per Fr. by Merulius Haller (1742), considering Von Haller’s first species (= Agaricus chantarellus L.) as type. He emended the genus so as to equal Cantharellus as compiled by Saccardo (Syll. Fung. 5: 482. 1887). Besides Von Haller’s two original species he listed Saccardo’s in an alphabetical enumeration. The first of these is Cantharellus albidus Fr., apparently sufficient reason for W. B. Cooke (Gen. Homobas. 62. 1953) to list it as type of Merulius “Hall, ex O. Kuntze”! — MONADELPHOUS HOMONYMS: (i) Merulius Fr., Syst. mycol. 1: 326. Jan. 1, 1821. — Type species (selected): Merulius tremellosus Schrad. per Fr.—See page 11. — (ii) Merulius S. F. Gray, Nat. Arrang. Brit. Pl. 1: 636. Nov.? 1821. —Type species (only species) : Merulius aurantiacus (Wulf.) Pers. (Agaricus aurantiacus Wulf.). — Gray misapplied the name Merulius [Haller] Boehmer (for which he cited no author) by excluding its type species (Agaricus chantarellus L.), his only (British) species being the one here indicated as type. Typonym: Hygrophoropsis (J. Schroet.) Maire apud Martin-Sans (1929). —TYPONYMS: Cantha-

rellus Adans. per Fr. (Jan. 1, 1821), Alectorolophoides Earle (1909). —STATUS. Impriorable on account of the earlier name Merulius Fr. As a later monadelphous homonym preferably to be treated as if it were a mere application of that name.

Merulioporia Bondarz. & Sing, apud Bondarz. in Sovietsk. Bot. 1943 (1) : 38 (n.v.). — ETYMOLOGY: the genus Merulius; the genus Poria. Gender: f. — TYPE SPECIES: Poria taxicola

(Pers.) Bres.—This species is often (apparently


incorrectly) identified with Polyporus rufus Schrad. per Fr. = Poria rufa (Schrad. per Fr.) Cooke. It is also called Polyporus haematodus Rostk. — PROTONYM: Merulioporia Bondarz. & Sing, in Ann. mycol. 39: 48. 1941.— Introduced for a group of species, of which two were mentioned by name, Poria taxicola (Pers.) Bres. being indicated as type species. — The generic name was not

validly published on this occasion ; no Latin description, no proper reference. The same applies to the following uses by the same authors: apud Singer (in Mycologia 36: 67, 69. 1944), apud Bondarzev (Trutov. Griby 51, 593. 1953). The authors refer to a previous publication (in Sovietsk. Bot.) where the name may have been validly published : I have not consulted this paper. — HOMONYM : Meruliporia Murrill (1942; see below).

Meruliporia Murrill in Mycologia 34: 596. 1942. — ETYMOLOGY: the genus Merulius; the genus Poria. Gender: f. —TYPE·SPECIES (only original species): Poria incrassata (Berk. & C.) Burt. — HOMONYM: Merulioporia Bondarz. & Sing, apud Bondarz. (1943; see above).

Mycodendron Mass. in J. Bot., Lond. 29: 1. 1891. — ETYMOLOGY: μυκής, fungus; δένδρον, 'tree.

Gender: n. — TYPE SPECIES (only original species) : Mycodendron paradoxum Mass.—A very

dubious fungus. The insinuation has been made that it never really existed except in Massee’s

imagination— perhaps justified. There is no type at Kew. —VARIANT SPELLINGS: “Mycodendrom”;

Léon March., Énum. méth. Mycoph. 203.1896.—An error. — “Mycodendrum”; Clem. & Shear,

Gen. Fungi 348. 1931.—A ‘correction.'

Mycodendrum.—See Mycodendron.

Serpula (Pers.) per S. F. Gray, Nat. Arrang. Brit. PI. 1: 637. 1821. — ETYMOLOGY: serpo, I creep. Gender: f. — TYPE SPECIES (selected): Merulius destruens Pers. = Merulius lacrimans (Wulf.) per Fr. —DEVALIDATED BASINYM: Merulius sect. Serpula Pers., Syn. Fung. 496.1801—Of the four species included the first is Merulius destruens Pers., undoubtedly to be selected as type species; one of others is Merulius serpens Tode, here mentioned only because its specific epithet might have inspired the sectional one. — Later Persoon (Traité Champ, comest. 43. 1818) spoke of this group as a genus: “Xylomyzon ou Serpula.” However, no description or reference was added and Serpula was thus published by Persoon, as a generic name, only as a nomen nudum;

compare quotation under Xylomyzon. Eventually he preferred Xylomyzon Pers. (1825), q.v., for the genus. — SCOPE. When Gray validly re-published the generic name, his only (British) species was Merulius destruens. —SYNISONYMS. Both Xylophagus Link per Murrill, q.v., and Xylomyzon Pers., q.v., have the same (devalidated) basinym as Serpula. —TYPIFICATION. Murrill (in J. Mycol. 9: 90, 101. 1903) considered Gray’s only species as type of “Serpula Gray.” Later authors who listed a type have accepted that species, for instance W. B. Cooke (in Mycologia 49: 201. 1957), who listed it as “S. destruens Pers. ex S. F. Gray.” — REMARK. “Serpula (Merulius *** Serpula Pers. Syn. p. 496) Karst. (N. gen.)” in Medd. Soc. Fauna Fl. fenn. 11: 20.1884 is an independent, re-publication of the generic name, for coloured-spored species (“Sporae plus minus late

ellipsoideae,


ferrugineae vel cinnamomeae”). First species: “Merulius lacrymans (Wulf.) Schum.”; the last two of seven examples mentioned are "M. aureus Fr. et M. molluscus Fr.” and these possess quite different spores. Afterwards Karsten (in Bidr. Känn. Finl. Nat. Folk 48. 1889) divided his genus into two, Serpula (p. 343) with M. aureus and M. molluscus and four other species, characterized

by ‘‘Sporerna mer eller mindre bredt elliptiska, hvita eller gulaktiga,” and Gyrophana Pat. (p. 345) with M. lacrimans and M. squalidus Fr. and characterized by “Sporerna rostfärgade.” In view of the original contents and description this latter use is a misapplication rather than a valid emendation. W. B. Cooke (Gen. Homobas. 89. 1953) listed Serpula “(Pers.) Karst.” with Merulius lacrimans Pers. as type species. —TYPONYMS: Xylomyzon Pers. (1825), Gyrophora Pat. (1887; preoccupied), Gyrophana Pat. (1897), and Xylophagus Link per Murrill (1903).

Sesia Adans. per Ο. Κ., Rev. Gen. PL 2: 869. 1891.—See Polyporaceae. —· When revalidating this name, O. Kuntze misapplied it by substituting it for Merulius Fr. as he found that genus treated by Saccardo (Syll. Fung. 6: 411. 1888). The type species is not identifiable with Merulius lacrimans (Wulf.) per Fr., as Kuntze thought, but with Lenzites sepiaria (Wulf, per Fr.) Fr.

Trabecularia Bonord. in Bot. Ztg 15: 211. 1857. — ETYMOLOGY: trabecula, small beam. Gender: f. —TYPE SPECIES (only original species): Trabecularia villosa Bonord.—Judging from description and figures this is Merulius tremellosus Schrad. per Fr. —TYPONYM: Merulius Fr. (1821).

Xylomyzon Pers., Mycol. europ. 2: 26. 1825. —ETYMOLOGY: ξύλον, wood; μύκης, fungus. Gender: m. — TYPE SPECIES (selected): Xylomyzon destruens (Pers. per S. F. Gray) Pers. = Merulius lacrimans (Wulf.) per Fr. —PROTONYM: Xylomyzon Pers., Traité Champ, com. 43. 1818.—A nomen nudum: “... les Cantharelloidées ... les genres dont elle est composé sont les Merulius (Cantharellus), Xylomyzon ou Serpula, Gomuphus [= Gomphus] et Daedalea”—nothing else. — BASINYM. See under “Remark.” — SCOPE. Introduced for about the same group to which Fries applied the name Merulius Fr. 1821. (Persoon used Merulius for Cantharellus Adans. per Fr.) — REMARK. From Persoon’s own remark of 1818 quoted above, one may conclude (entirely justifiably I believe) that Xylomyzon and Serpula are two names for the same taxon.

Indeed, Xylomyzon, as a generic name, covers exactly the same group as was previously called Merulius sect. Serpula Pers. 1801; see under Serpula (Pers.) per S. F. Gray. — TYPIFICATION.

Persoon’s first species is a well known destructive house fungus and it may be concluded that it was his leading species. It was already adopted as type species by Murrill (in J. Mycol. 9: 91, 102. 1903) and W. B. Cooke (Gen. Homobas. 100. 1953). —VARIANT SPELLING: “Xylomycon” ; Reichenb., Consp. Regni veg. 14. 1828 (as a synonym). —TYPONYMS: Serpula (Pers.) per S. F. Gray (1821), Gyrophora Pat. (1887; preoccupied), Gyrophana Pat. (1897), and Xylophagus Link

per Murrill (1903).

Xylophagus Link per Murrill in Torreya 3: 7.1903 ; in J. Mycol. 9: 90,102.1903. —

ETYMOLOGY: ξύλον, wood; φάγος, glutton. Gender: m. —- TYPE SPECIES


(selected) : Merulius destruens Pers. = M. lacrimans (Wulf.) per Fr. — DEVALIDATED BASINYM:

Merulius sect. Serpula Pers., Syn. Fung. 496. 1801.— See for this taxon under Serpula (Pers.) per S. F. Gray. — DEVALIDATED GENERIC NAME: Xylophagus Link in Mag. Ges. naturf. Fr. Berl. 3: 38. 1809.— “Hujus loci Merulii familia Serpula a Persoonio dicta.” —VALID PUBLICATION. As far

as I am aware this name was not validly published before it was deliberately taken up by Murrill. It was once listed after 1821 as “Xylophagus” (name only) by Link (in Abh. phys. Kl Akad. Wiss. Berlin 1824: 179. 1826), without a reference (not even an author’s citation) or description however. — The following is Murrill’s first use (in full) when he definitely accepted the generic

name:

“A new family of the Basidiomycetes.—Xylophagaceae. This family is based on Xylophagus Link Berl. Mag.

3: 38. 1809. (Merulius Hall.) and allied genera formerly included in the Polyporaceae. Its distinguishing

character is a gelatinous and at the same time porous hymenium. Its genera may be grouped under three

subfamilies: The Favolaschiae, including plants wholly gelatinous both in context and hymenium, the

Xylophageae, in which the context varies from semi-gelatinous to firm or fibrous and the Gloeoporeae, with

firm fibrous context and a hymenium of deep cohering tubes instead of shallow reticulations. This last group

approaches the Polyporaceae, but differs by reason of its separable gelatinous hymenium. The types of the

three subfamilies mentioned above are Favolaschia Pat., Xylophagus Link and Gloeoporus Mont.”—Murrill (in

Torreya 3: 7. 1903).

As one will notice, Murrill did not give a description but only a reference to Link’s (now) devalidated name; the latter itself was not accompanied by a description either, but Link referred to its basinym, which at last was accompanied by a description. — SCOPE. “In 1825, Persoon proposed the name Xylomyzon for the same group”—Murrill (in J. Mycol. 9: 90. 1903). — REMARK. In view of the references, Xylomyzon is to be taken as a straight isonym of Merulius

sect. Serpula Pers. and the same species that has been selected for that sectional name has to be accepted as type of Xylophagus Link, per Murrill, too. This is Merulius destruens (see under Serpula) and it was already indicated by Murrill (in J. Mycol. 9: 90, 102. 1903) as such under the

name of “X. lacrymans (Wulf.)”:

“The name was proposed for the section of Merulius called Serpula by Persoon in his Synopsis, 496, 1801 and

is therefore to be considered as based upon X. lacrymans (Wulf.) and the four other species listed there.”—

Murrill (in J. Mycol. 9: 90. 1903).

VARIANT SPELLING: “Xylophflgus”; Ainsw. & Bisby, Diet. Fungi, 2d Ed., 374. 1945.—A printing-

error. — TYPONYMS & SYNISONYMS: Serpula (Pers.) per S. F. Gray (1821), Xylomyzon Pers. (1825),

Gyrophora Pat. (1887; preoccupied), and Gyrophana Pat. (1897).

NOTES ON RESUPINATE HYMENOMYCETES—V*

M. A. Donk


The genera formerly included in Botryobasidium Donk and Ceratobasidium D. P. Rog. are reviewed. They are, in the order of the discussion, Ceratobasidium, Oliveonia Donk, Uthatobasidium Donk, Botryohypochnus Donk, Botryobasidium, Thanatephorus Donk, and Koleroga Donk.

Oliveonia is a new name for Heteromyces L. Olive (preoccupied), while Koleroga is the name of a new genus based on the kole-roga fungus. Uthatobasidium sect. Tpsilonidium Donk is a new section based on Corticium sterigmaticum Bourd. Botryohypochnus and Botryobasidium are re-defined. New specific combinations are made with Oliveonia (2), Uthatobasidium (2), and Botryobasidium (1). Koleroga noxia Donk, the true kole-roga fungus, is described as a new species.

INTRODUCTION.—The genus Botryobasidium Donk (1931: 117) as originally conceived corresponded exactly with Corticium sect. Botrydea Bourd. & G. It was soon emended by Rogers (1935: 10) by the exclusion of some species which were placed in Ceratobasidium D. P. Rog. ; and

by the inclusion of Botryohypochnus Donk (Tomentella subsect. Botrytes Bourd. & G.). Still later Rogers (1943: 95) introduced the kole-roga fungus into the genus and in connection with this transfer felt bound to replace the generic name Botryobasidium by the earlier one Pellicularia Cooke. This latter name was rejected by Donk (1954) who pointed out that it was illegitimate (not available). Moreover, it is my opinion that the kole-roga fungus is not at all congeneric with the typical species of Botryobasidium. On the earlier occasion I did not formally propose a new generic name for the kole-roga fungus, because no good type collection was available. This situation has not improved, but none the less a new genus will be introduced for it this time. A few years ago Donk (1936a) also excluded the species with spores exhibiting repetition as two small distinct genera, Uthatobasidium Donk and Thanatephorus Donk. This character of the spores was considered as of generic value also by Pilât (1957: 80) and Olive (1957: 431), but instead of keeping the species apart, these authors transferred them to Ceratobasidium which agrees in the spores exhibiting repetition. This would make Ceratobasidium still more heterogeneous than it already is and these transfers will be questioned below.

Recently Eriksson ( 1938a : 2 ; 1958c : 18) again separated Botryohypochnus from

Botryobasidium and, in addition (1958b: 58; 1958c: 19), showed himself in favour of the maintenance of both Uthatobasidium and Thanatephorus. Finally, Eriksson (1958a : 3 ; 1958b : 47) divided the remnants of Botryobasidium into three subgenera. The separation of Botryohypochnus he based on the shape and spines of the spores. A suggestion will be made below to emend the genus, primarily on the basis of basidial morphology.

A discussion and survey of the genera formerly included in Botryobasidium follows. I have also entered Heteromyces L. Olive although it was recently

* Part I was published in Reinwardtia 2: 425-434. 1954; Part II, in Reinwardtia 3: 363-379· 1956; Part III, in Fungus 26: 3-24. 1956; Part IV, in Fungus 27: 1-29. 1957.

Nov. 1958] M. A. Donk: Notes on Resupinate Hymenomycetes—V 17

proposed for a segregate of Ceratobasidium rather than of Botryobasidium. Since the name is preoccupied it is replaced by the isonym Oliveonia Donk.

KEY TO THE GENERA FORMERLY INCLUDED IN BOTRYOBASIDIUM AND CERATOBASIDIUM

I . Spores exhibiting repetition. Sterigmata stout, often somewhat fusoidly swollen when starting to produce spores, usually straight, but may be horn-shaped (i.e. curved outwards) in Oliveonia, 2-4(-5) per basidium. Conidiferous stages lacking. 2. Fruit-body not typically Botryobasidium-like in structure, in some species continuous, mucous-gelatinous,

somewhat fleshy, or waxy, in others delicate, pruinose (somewhat waxy when fresh), or a tender weft. Basidia in the more typical species1 distinctly swollen (about 2-3 x as wide as supporting hyphae), many abruptly narrowed towards bases and distinctly (although often short-) stalked above basal septa (Tulasnella-like). Hyphae usually rather thin, 2-6(-9) μ wide (not counting inflations) ; clamps lacking or present.

3. Thin-walled cystidia lacking. Saprobic, or (in one species) parasitic and forming sclerotia-like bodies. Ceratobasidium 3. Thin-walled cystidia (gloeocystidia) present. Saprobic. Oliveonia 2. Fruit-body Botryobasidium-Yike: floccose, under the lens granulose by separate clusters of basidia borne on

stout, erect, branched, short-celled, ascending hyphae (which rarely may be reduced in some parasitic forms) ; neither mucous-gelatinous, fleshy, or waxy, nor continuous. Basidia plump and short, hardly swollen, their bases never constricted or substipitate, broadly attached (as in Botryohypochnus isabellinus). Hyphae (rather) stout, 6-12 μ wide; clamps lacking. Gloeocystidia lacking.

4. Saprobic; no Rhizoctonia state. Uthatobasidium 4. Parasitic ; mycelium often in part soil-harboured, also completely aerial, developing into a Rhizoctonia

state: tending to grow in fibrils or strands and usually forming sclerotia-like bodies. Thanatephorus I. Spores not exhibiting repetition. Sterigmata tiny to rather stout, horn-shaped (i.e. curved outwards). Conidiferous stages may be present.

5. Saprobic: not developing into thread blights on trees or shrubs. Basidia formed in notable clusters on well developed ascending, branched hyphae (the clusters causing the granulose appearance of the fruit-body when seen under the lens), broad at bases, not much broader than supporting hyphae; sterigmata small. 6. Basidia plump, somewhat barrel-shaped, subobovoid, or subclavate, neither urnshaped nor

constricted at middle; sterigmata (2-)4· Botryohypochnus 6. Basidia rather short and plump to tubular, more or less distinctly constricted at middle or rather urn-

shaped; sterigmata 4-8 (usually 6), except perhaps in one species where they may number 4. Botryobasidium

5. Parasitic: developing into thread blights on trees or shrubs, the fruit-bodies usually formed on the underside of living leaves. Basidia on short, lateral branches of the repent hyphae of the fruit-body, either solitary or a few together, when young globular and swollen (much broader than supporting hyphae), often short-stipitate above basal septa; sterigmata rather stout, 2-4(-6) per basidium. Koleroga

CERATOBASIDIUM D. P. Rog.

Ceratobasidium D. P. Rog. in Stud. nat. Hist. Univ. Iowa 17 (1) : 4. 1935.

This genus was published with four species, Ceratobasidium calosporum D. P. Rog. (type species), Corticium cornigerum Bourd., Ceratobasidium obscurum D. P. Rog., and Corticium sterigmaticum Bourd., all well described and illustrated with figures of microscopical details. The two species previously published by Bourdot were taken from Corticium sect. Botryodea Bourd. & G., i.e. Botryobasidium Donk.

The differences from Botryobasidium were not clearly stated and, as indicated

1 In Ceratobasidium atratum (Bres.) D. P. Rog. apud G. W. Mart, the basidia seem to vary considerably and have been likened to a hydra.


in a previous paper (Donk, 1956a: 370), mainly of an illusory nature: the sterigmata were interpreted as ‘epibasidia’, thus as something quite different, but how exactly these ‘epibasidia’ were to be distinguished from sterigmata was not explained. Stout ‘epibasidia’ (sterigmata) and repetition of the spores, two of the characters that played an important rôle in the generic

description are also typical of those species of Botryobasidium that have been referred to the genera Uthatobasidium Donk and Thanatephorus Donk. Rogers conceded that ‘ ‘the structure and texture of the fructification in [Corticium] sterigmaticum differ in no important respect from those characterizing Botryobasidium, to which truly it is closely allied.” This species may be excluded from Ceratobasidium and is perhaps best referred to Uthatobasidium (cf. p. 21).

Yet the first three of the original species and a few later additions do seem worthy of separation from Uthatobasidium, which is also saprobic and shares in addition the stout sterigmata and the repetition of the spores. This does not mean that all the species now combined in Ceratobasidium really form a homogeneous taxon.

Of the original species two may be opposed to the rest, viz. C. calosporum and C. cornigerum. The features that may be emphasized for their differentiation from Uthatobasidium are the different, not typically Botryobasidium-like, structure of the fruit-body, which is, moreover, tenuous, pruinose, somewhat waxy when fresh; the diameter of the hyphae which is inferior to that in Uthatobasidium, about 3-6(-7·5)μ wide; and the broad and swollen basidia (if compared with the supporting hyphae), about 10 μ wide, and (at least many of them) more or less

substipitate above the basal septa, i.e. Tulasnella-like. As thus defined this kernel of saprobic species should perhaps also include the parasitic Tulasnella anceps Bres. & H. Syd. apud H. Syd. This latter species differs not only in being parasitic, but also in forming a kind of Rhizoctonia- like ‘sclerotia’, and in having a more arid fruit-body, characters that might suggest its assignment to Thanatephorus.

Jackson (1949: 243) once remarked, on the occasion of the transfer of Tulasnella anceps to Ceratobasidium, that the latter genus and Pellicularia Cooke sensu D. P. Rog. (= Botryobasidium s. lat.) approach each other very closely, especially through the parasitic species. He concluded that Tulasnella anceps belonged to Ceratobasidium “because of the character of the basidium [showing the division into hypo-basidium and epibasidia] and the habit of spore germination by repetition.“ He could easily have replaced ‘Ceratobasidium’ by ‘Botryobasidium’, because both characters he emphasized were also to be found in certain species of the latter genus (like Hypochnus solani) as conceived at that time. It is likely that the real reasons why he decided in favour of Ceratobasidium, and correctly so I would say, were the

hyphae, narrow (3.5-5.5 μ wide) in comparison with the basidia, which appear as strongly swollen bodies ; the shape of the basidia, many of which are slender at their bases and more or less substipitate above the basal septa; and the poor development of the fertile branches which arise from the subicular hyphae to form the clusters of basidia so characteristic in typical species of the Botryobasidium group. The basidia may even arise directly from the hyphae that form the delicate subicular weft of the fruit-body. In all these regards Tulasnella anceps closely resembles the two typical species of Ceratobasidium (especially C. cornigerum).


Rogers’s third original species, C. obscurum, according to its description, differs widely in forming mucous-gelatinous continuous layers, and in having the constituent elements of the fruit-body embedded. It would seem that in admitting this species, Ceratobasidium decidedly becomes an artificial taxon. In such a genus a few more aberrant species like C. atratum (Bres.)

D. P. Rog. (apud Martin, 1941: 262), attributed to it by Martin (1939: 513 fs. 21-27, as C. plumbeum G. W. Mart.), will do no harm either for the time being. This fungus forms waxy to somewhat fleshy, continuous fruit-bodies with a definite palisade-like hymenium. Ceratobasidium atratum has been made the type of a new generic name, Hydrabasidium Parker-Rhodes (1954: 325, 338; name not validly published; cf. Donk 1957: 73), and one might consider taking it up, but this solution would be only a partial one. Ceratobasidium in its present circumscription as a whole is in need of a critical study.2 As recently described by Wakefield (1952: 64 f. 36) the basidia of C. atratum may be irregular and Hydra-like in appearance. For the group of cystidiate species which Donk temporarily put into Ceratobasidium and which Olive has placed in a genus of their own (Heteromyces), see Oliveonia.

Recently Wakefield (1952: 64 f. 37) transferred Corticium terrigenum Bres. to Ceratobasidium, although no repetition of spores was found. She described the fruit-body as continuous and somewhat fleshy (it has a definite hymenium), with four stout sterigmata per basidium. In certain important respects this species reminds one of Cerinomyces G. W. Mart. (cf. Donk, 1936a: 375)3 and may be a four-spored member of that genus. It is doubtful whether it is

to be admitted to the present genus. On the whole certain species of Ceratobasidium remind one in too many respects of

Tulasnella as to leave much room for doubt that they are closely related with the latter genus: the fruit-bodies of some of its species have the same appearance and internal structure, and the characters emphasized above for the hyphae and the basidial bodies as well as their relative dimensions are the same. The one and only really diagnostic difference given in literature is in the sterigmata: these are strongly developed in Ceratobasidium, but are never as inflated and voluminous as in Tulasnella; they are not typically separated from the basidial body by walls across their bases.4 On the other hand the relation of Ceratobasidium with Uthatobasidium, q.v., might also be close, and differences of opinion have arisen (as already briefly indicated above) as

to whether certain species are to be referred to this or to that genus (see p. 21).

2 Parker-Rhodes (1954: 335) states that, “The genus Ceratobasidium Rogers was regrettably founded on C. calosporum Rog. as type, and as this appears to be only a form of Prototremella calospora Boud. with aseptate basidia, the name Ceratobasidium cannot stand, though it is certainly a good genus.” Since Prototremella calospora [= Tulasnella calospora (Boud.) Juel] has aseptate basidial bodies and quite different sterigmata it may be that Parker-Rhodes confuses this species with Sebacina calospora Bourd. & G. It is a pity that he does not explain more fully his somewhat startling conclusion.

3 The suggestion made by Eriksson (1958b: 46) and supported by Donk (1956a: 375) that Cerinomyces would not belong to the Dacrymycetaceae but rather to the Corticiaceae has been declared “utterly fantastic and completely without merit” by Martin (1957: 25) who also promised to return to this question. In the meantime a discussion from this side will be postponed.

4 Martin (1937: 25) does not want to call the Tulasnella basidium holobasidious on account of these cross-walls. In my opinion there is no reason to follow him in this respect as I have already pointed out recently (Donk, 1958: 96-98).


The slender and remarkable spores of C. calosporum recall to mind those of Sebacina calospora Bourd. & G. and Gloeotulasnella calospora (Bourd.) D. P. Rog.; the resemblance is not restricted to this feature and seems of a more general nature except for the basidia. This might be taken as indicating a close relationship between the three genera involved.

Summarizing, Ceratobasidium is maintained for the time being in a wide circumscription as an artificial genus, for want of a better solution. It may be characterized thus:—

Saprobic; or, in one species, parasitic and developing appressorial pads or infection cushions, but not a

typical Rhizoctonia state. Fruit-body strictly resupinate, effused, indeterminate, delicate and consisting of a

(very thin) weft or film, or pruinose, to continuous, arid or waxy when fresh to somewhat fleshy or mucous-

gelatinous; structure not typically Botryobasidium-like. Cystidia or gloeocystidia lacking. Basidia in the more

‘arid’ and typical species strongly inflated and (at least many) constricted at bases and thus appearing

substipitate above basal septa, much wider than supporting hyphae which are rather narrow, 3 -6(- 7.5) μ in

diameter. Sterigmata usually stout, but neither as strongly inflated as in Tulasnella (and Gloeotulasnella) nor (or

exceptionally) separated from basidial body by walls across their bases. Spores exhibiting repetition. Conidial

stages not reported.

TYPE SPECIES (by original designation).—Ceratobasidium calosporum D. P. Rog.

EXAMPLES.—

1. Fruit-body delicate, pruinose, or a very thin weft or film, arid to only slightly waxy. No continuous hymenium. Basidia strongly inflated, mostly substipitate above basal septa. 2. Saprobic.—Ceratobasidium calosporum D. P. Rog., C. cornigerum (Bourd.) D. P. Rog. 2. Parasitic, developing sclerotia-like bodies.—Ceratobasidium anceps (Bres. & H. Syd. apud

H. Syd.) H. S. Jacks. 1. Fruit-body mucous-gelatinous, waxy, or somewhat fleshy, continuous.—Ceratobasidium obscurum

D. P. Rog., C. atratum (Bres.) D. P. Rog. apud G. W. Mart.

Oliveonia Donk, nom. nov.

Heteromyces L. Olive in Amer. J. Bot. 44: 432. 1957, basinym; not Heteromyces Müll.-Arg. in Flora 72: 505. 1889 (Lichenes, Cladoniaceae).

TYPE SPECIES (by original designation).—Sebacina fibrillosa Burt.

Examples.—Oliveonia fibrillosa (Burt) Donk, comb. nov. (basinym, Sebacina fibrillosa Burt in Ann. Missouri

bot. Gdn 13: 335. 1926)5 and Oliveonia pauxilla (H. S. Jacks.) Donk, comb. nov. (basinym, Corticium pauxillum

H. S. Jacks, in Canad. J. Res. C 28: 724. 1950)·

The transfer to Ceratobasidium of the gloeocystidiate species Sebacina fibrillosa Burt (originally erroneously described by its author as having cruciate-septate basidia) by Rogers & Jackson (1943: 327) and Martin (1948: 113 f. 1) raised the question, why Peniophora heterobasidioides (Rogers, 1935 : 30 f. 5) and Co.rticium pauxillum (Jackson, 1930: 724 f. 9)6 should not follow. Both are also gloeocystidiate and also exhibit repetition of the spores. Their sterigmata do not differ essentially from the ‘epibasidia’ of C. fibrillosum (as drawn by Martin).

These species were placed together in Donk’s group (vi) of the “tulasnelloid fungi”, which he temporarily included in the already heterogeneous genus

5 Olive (1957: 432) regards Peniophora heterobasidioides D. P. Rog. as a synonym of Sebacina fibrillosa.

6 The author of Corticium pauxillum placed it in an emended Corticium sect. Athele Bourd. & G., but it was indicated that possibly it had to be included in Ceratobasidium.


Ceratobasidium (1956: 373, 377). Olive raised it to generic rank and called it Heteromyces, a name which is preoccupied and, therefore, to be replaced by a new one.

A simultaneously described genus is Metabourdotia L. Olive (1957: 429), also possessing gloeocystidia. It differs in its basidia which are often incompletely septate in a tremellaceous

manner. It seems best referred to the Tremellaceae and would seem to differ hardly, if at all, from certain members of Sebacina Tul. as currently emended in agreement with McGuire, that

is, in a wide sense, inclusive of gloeocystidiate species.

UTHATOBASIDIUM Donk

Uthatobasidium Donk in Reinwardtia 3: 376. 1956. Saprobic; not forming Rhizoctonia states. Like Botryobasidium (sensu stricto, see p. 26) in consistency and

structure of fruit-body, but spores exhibiting repetition and with rather large, truncate apiculus; sterigmata

stout, usually somewhat fusoidly swollen when starting to produce spores, straight rather than curved

outwards ; basidia neither constricted at middle nor urn-shaped, with 2-4 sterigmata. Conidial stages not

reported.

TYPE SPECIES.—Hypochnus fusisporus J. Schroet. Genotype distribution: Fuck., Fung, rhenan. exs. No. 2396, as

Hypochnus flavescens Bonord.

EXAMPLES.—

I. Sterigmata (2—)4, never constantly 2, shorter than the basidial body, fusoidly swollen. Sect. Uthatobasidium.—Uthatobasidium fusisporum (J. Schroet.) Donk, see below; U. ochraceum (Mass.) Donk, see below.

I. Sterigmata 2, very stout, about as long as the basidial body, attenuate-cylindrical, divergent. Uthatobasidium sect. Ypsilonidium Donk, sect. nov?—Type species, Corticium sterigmaticum Bourd.

Uthatobasidium is a segregate from Botryobasidium Donk (Pellicularia Cooke sensu D. P. Rog.). It seems more closely related to Botryohypochnus Donk rather than to Botryobasidium on account of the characters of the basidia. The differences from Botryobasidium are given in the above generic description. It is also closely related to Thanatephorus Donk, but the species of that genus are parasitic and the mycelium develops typical Rhizoctonia states, while also the ascending basidiferous hyphae may be less typically Botryobasidium-like branched or, even, may be poorly developed.

Uthatobasidium can readily be distinguished from Ceratobasidium by its basidia, which are not Tulasnella-like (not strongly inflated and never constricted at their broad bases) ; by the wider and firmer hyphae ; as well as by their invariably typically Botryobasidium-like structure with ascending, branched hyphae bearing the clusters of basidia.

I can see no reason to refer Corticium sterigmaticum to Ceratobasidium as was done by

Rogers (2935: 7; see also the present paper, p. 18). The structure of its fruit-body (well developed, branched, ascending hyphae and clusters of basidia), dimensions of the hyphae, the shape of the basidial body, and other features are all those of Uthatobasidium. It differs from the typical species of the latter genus mainly in its sterigmata: these are very strongly developed (hence ‘epibasidia’ to Rogers) and invariably two in number, features sufficient it would seem to place it in a special section.

7 Sterigmata 2, attenuato-cylindrica, divergentia, basidia plus minusve aequilonga.


Uthatobasidium fusisporum (J. Schroet.) Donk, comb. nov.

Hypochnus fusisporusj. Schroet. in Krypt.-Fl. Schles. 3 (1): 416. 1888, basinym of new combination. —Corticium fusisporum (J. Schroet.) W. Brinkm., Westf. Pilze No. 53. 1904, misapplied; not Corticium fusisporum Cooke & Ell. in Grevillea 8: 11. 1879. — Peniophora fusispora (J. Schroet.) Höhn. & L. in Ann. mycol. 4: 289. 1906, misapplied.

Coniophora vaga Burt in Ann. Missouri bot. Gdn 4: 251 f. 8. 1917. —Corticium fenestratum Overh. in Mycologia 26: 510 pl. 55 f. 5. 1934 (“nom. nov.”), isonym. — Holotype: U.S.A., New York, Hudson Falls (S. H. Burnham 20, MO 54498). — According to D. P. Rog. in Farlowia 1: 105, 107. 1943 = Pellicularia flavescens (Bonord.) D. P. Rog. [sensu Fuck.].

MISAPPLICATIONS.—Hypochnus flavescens Bonord., Handb. Mykol. 160. 1851 sensu Fuck, in Jb. nassau. Ver. Naturk. 25-26: 291. 1871; Wint. in Rabenh. Krypt.-Fl., 2. Aufl., 1 (1): 329. 1882 (Corticium), in part; D. P. Rog. in Stud. nat. Hist. Univ. Iowa 17 (1): 13 f. 8. 1935 (Botryobasidium); D. P. Rog. in Farlowia 1: 105. 1943 (Pellicularia), in part; L. Olive in Amer. J. Bot. 44: 431. 1957 (Ceratobasidium).

Descriptions & illustrations.—Von Höhnel in Oesterr. bot. Z. 54: 428. 1904 (Corticium flavescens; in obs. to C.

viride) ; Wakefield & Pearson in Trans. Brit, mycol. Soc. 6: 317 fig. 1920 (Corticium flavescens); Bourdot & Galzin,

Hym. France 239. 1928 (Corticium flavescens); Rogers in Stud. nat. Hist. Univ. Iowa 17 (1): 13 f. 8. 1935 (Bo-

tryobasidium flavescens) ; Eriksson in Symb. bot. upsal. 16 (1) : 59 f. 12a-e. 1958 (Botryobasidium flavescens).

TYPE LOCALITY.—Silesia, Neumarkt, Lissa.

This species has been identified by modern authors with Hypochnus flavescens Bonord. in imitation of Fuckel (l.c.), whose distribution of the fungus he so identified authenticates his interpretation (Fuckel, Fung, rhenan. exsic. No. 2396. 1871). This identification is extremely doubtful, to say the least. Of Bonorden’s type nothing is known but its description, which runs:—

“Hypochnus flavescens m., hat ein zartes, ausgegossenes Stratum von körnigem Ansehen und Weissgrauer Farbe, wird später gelb. Die konischen Basidien tragen 4 rundliche Sporen an langen Stielen.”—Bonorden (l.c.).

To me Hypochnus flavescens Bonord. is a nomen dubium, the more so as one would have to

allow for an erroneous spore-shape if identifying it with the present species; it might conceivingly be the next species, Uthatobasidium ochraceum (Mass.) Donk but such an identification, too, is far from certain.

The identity of Hypochnus fusisporus J. Schroet. was established by Rogers (1943: 103) by a careful analysis of the original description. This fungus has been confused by Von Höhnel & Litschauer and subsequent authors with one Rogers identified with Coniophora ochroleuca Bres. apud W. Brinkm.

The spores of U. fusisporum are highly characteristic, consisting if it were of a globular body with two broad opposed gibbous outgrowths (of which one bears the apiculus) so that the spore appears lemon- or spindle-shaped; the protuberances may vary in development, but seem constantly present. When in a preparation a number of the spores may appear globular this is often due to their position. “En faisant voyager les spores par une légère pression de la lamelle, on se rend compte que les spores sphériques présentent deux gibbosités ou mamelons plus ou moins accusés, dans le prolongement d’un axe excentrique.”—Bourdot & Galzin (1928: 239).

Rogers (1943: 105) after first having distinguished between the present species and

Botryobasidium ochraceum (Mass.) Donk apud D. P. Rog. = Uthatobasidium ochraceum (Mass.) Donk., the next species, changed his mind

Nov. 1958] M. A, Donk: Notes on Resupinate Hymenomycetes—V 23

and afterwards combined the two. He based his conclusion on the variation occurring in the development of the protuberances of the spores in U. fusisporum and stated that there are intermediates between the types of spores. It may be that these occur, but then in individual spores of preparations of U. fusisporum showing a majority of typical spores. The reverse, spores

developing protuberances in U. ochraceum, I have not encountered and this is suggestive.

Eriksson (1958b: 58) also concluded that there are good reasons to keep the two species apart.

Uthatobasidium ochraceum (Mass.) Donk, comb. nov.

? Hypochnus flavescens Bonord., Handb. allg. Mykol. 160. 1851 (nomen dubium).—Corticium flavescens (Bonord.) Wint. in Rabenh., Krypt.-Fl., 2. Aufl., 1 (1) : 329. 1882, in part; Mass. in J. Linn. Soc., Lond. (Bot.) 27: 149. 1890, misapplied ; not Corticium flavescens Bres. in Ann. mycol. 3: 163. 1905. — Botryobasidium flavescens (Bonord.) D. P. Rog. in Stud. nat. Hist. Univ. Iowa 17 (1): 13 f. 8. 1935, misapplied. — Pellicularia flavescens (Bonord.) D. P. Rog. in Farlowia 1: 105. 1943, misapplied at least in part? —Ceratobasidium flavescens L. Olive in Amer. J. Bot. 44: 431. 1957 (incomplete bibliographic reference to basinym), misapplied at least in part?

Coniophora ochracea Mass. in J. Linn. Soc., Lond. (Bot.) 25: 137 pl. 47 f. 13. 1889, basinym of new combination. — Botryobasidium ochraceum (Mass.) Donk apud D. P. Rog. in Stud. nat. Hist. Univ. Iowa 17 (1): 16 ƒ. 7. 1935.

Corticium frustulosum Bres. in Ann. mycol. 1: 98. 1903; in Ann. mycol. 9: 425. 1911. — Type: ‘Poland’ (B. Eichler, S). — According to John Erikss. in Symb. bot. upsal. 16 (1): 59. 1958 = Botryobasidium ochraceum (Mass.) Donk.

Corticium frustulosum var. intermedia Bourd. & G., Hym. France 240. “1927” [1928]. — Type locality: France.

DESCRIPTIONS & ILLUSTRATION.—Rogers in Stud. nat. Hist. Univ. Iowa 17 (1): 16 J· 7· 1935 (Botryobasidium

ochraceum); Bourdot & Galzin, Hym. France 239, 240. (Corticium frustulosum, C. frustulosum var. intermedia).

TYPE LOCALITY.—England, Kew. Type (K) apparently lost.

The description by Rogers (1935: 16 f. 7) was based on two collections, of which one (C. A. Brown 486) was found in North America. Afterwards he fused the globose-spored specimens with “Pellicularia flavescens”, the preceding species, which is here called U. fusisporum (J. Schroet.) Donk.

Bourdot & Galzin kept the fungus they identified with Coniophora ochraceum distinct from Corticium frustulosum Bres. as variety intermedia, indicating that it is “moins épais que C. frustulosum et jamais blanc.” The epithet ‘intermedia’ was apparently chosen in view of a supposed variation of the spores in the direction of their “Corticium flavescens” (= U. fusisporum):—

Corticium frustulosum var. intermedia “ ... diffère de C. flavescens par ses spores qui sont presque toujours globuleuses, avec mucron assez long. On trouve ... [des spores qui] présentent un petit mamelon peu marqué sur le côté de la spore, près du sommet. Ces spores émettent un filament germinatif.”—Bourdot & Galzin (1928: 240).

The lateral protuberance is, I presume, the initial stage of the sterigma which produces the secondary basidiospore rather than the apical wart discussed under U. fusisporum; in Bourdot & Galzin’s description of Corticium frustulosum no repetition of the spores is mentioned and was apparently overlooked and, therefore, thought to be absent. The French authors did notice repetition in V. fusisporum.

When Von Höhnel & Litschauer (1906 : 1607; 1907 : 835) redescribed the preceding species

under the name of Corticium flavescens Bonord. sensu Fuck.,


they cited exsiccati representing U. fusisporum, but it may be that they already included the present species. This might be an explanation of the globular spores they mentioned in addition to the spindle-shaped ones, perhaps a second and co-existing explanation beside the one already discussed under U. fusisporum. This would explain why they declared Corticium frustulosum

synonymous with C. flavescens after the study of ‘an original specimen’ (Von Höhnel & Litschauer, 1908: 1082). This conclusion was opposed by Bresadola (1911), who pointed out that the spores of his species were “fere globosa, vel obovate” a fact verified by Rogers (1943: 107) for the type.

I have studied neither the type of Corticium frustulosum Bres. nor that of Coniophora ochracea Mass., and tentatively follow Bourdot & Galzin in combining the two into a single species. Eriksson (1958b: 59 f. 12f) depicts an ellipsoid spore taken from the type collection of C. frustulosum.

It should be remarked that Massee said nothing about the sterigmata in his description, but

rendered them as thin, straight, and short and presumably drew them too schematically.

BOTRYOHYPOCHNUS Donk

Tomentella J.-Olsen apud Bref., Unters. Gesamtgeb. Mycol. 8: 9. “1889” [1888]; not Tomentella Pat., Hym. Europe 154. 1887; not Tomentella P.Karst. in Bidr. Känn. Finl. Nat. Folk 48: 419. 1889. — Type species (selected): Tomentella flava Bref., asto basidial portion = Corticium isabellinum (Fr. per Fr.) Fr. — Cf. Donk 1957: 118-119.

Tomentella sect. Tomentellastrum [subsect.] Botrytes Bourd. & G. in Bull. Soc. mycol. France 40: 137. 1924. — Type species (selected): Tomentella isabellina (Fr. per Fr.) Höhn. & L.

Botryohypochnus Donk in Meded. Nederl. mycol. Ver. 18-20: 118. 1931.

Differing from Botryobasidium Donk sensu stricto (see p. 26) in the basidia which are rather short and barrel-

shaped, subobovoid or subclavate, plump (neither more or less constricted at middle nor urn-shaped), with

rounded apex bearing (2-)4 rather stout sterigmata. Spores not exhibiting repetition; surface ornamented (or

smooth in one dubious species). Conidial stages belonging to Ostracoderma Fr. (Phymatotrichum Bonord.)

reported but presumably not belonging here.

Saprobic, on rotten wood.

TYPE SPECIES (by original designation).—Hypochnus isabellinus Fr.

EXAMPLES.—Botryohypochnus isabellinus (Fr. per Fr.) John Erikss. ; Tomentella granulata Bref, (see below);

Pellicularia biapiculata D. P. Rog. apud G. W. Mart, in Lloydia 7: 71 f. 6. 1944.

TOMENTELLA GRANULATA Bref.

Tomentella granulata Bref., Unters. Gesamtgeb. Mykol. 8: 11 pl. 1 f. 15. “1889” [1888]. — Hypochnus brefeldii Sacc., Syll. Fung. 9: 243. 1891, isonym. — Botryobasidium granulatum (Bref.) Donk in Meded. Nederl. mycol. Ver. 18-20: 118. 1931.

The following description is drawn up from Brefeld’s account of the fungus ; the characters

between brackets in the description are taken from his figures:—

Fruit-bodies a dense, extended, granular, white covering. Hyphae (“die Mycelien”) very coarse, thick

(without clamps, only slightly narrower than those of B. isabellinum, the basidia formed on ascending hyphae).

Basidia not united to a hymenium, (slightly clavate, rather short, but not as short as in B. isabellinum, about 2-

2.5 times as long as


wide); sterigmata 4 (see Brefeld, op. cit. p. 9), (slender-conical, not inflated, curved outwards, 1-1,5 times as

long as the diameter of still attached, and thus perhaps not fully matured, spores). Spores globular, smooth,

10 μ in diameter8 (those still attached to the basidia about 8.5 μ, only slightly smaller than those of B.

isabellinum) ; repetition not mentioned.

HABITAT.—On soil, rarely on trunks. [Germany.]

This fungus has not yet been redescribed or identified. Rogers (1943: 107) cited it as a doubtful synonym of “Pellicularia” pruinata (Bres.) D. P. Rog. ex Linder, as treated by Rogers a species-complex having a wide range in spore form: “subglobose or short-ellipsoid, to broad ellipsoid-fusiform, more or less flattened toward the lateral apiculus, distally obtuse, (3.5—)4—7 (—7.5) x 3-5(-6)μ”, and the number of sterigmata per basidium (5-)6(-8). In my opinion T. granulata is clearly different in its larger, more regular-globose spores, and four-spored basidia.

Another possibility to be taken into consideration is that T. granulata belongs to one of those globular-spored forms of Uthatobasidium Donk, in this paper included in U. ochraceum (Mass.) Donk, which has spores of about the size as indicated by Brefeld, but they exhibit repetition and the sterigmata, too, are quite different, neither slender-conical nor curved outwards, but usually stouter and more or less ventricosely swollen. There is no reason to assume incorrect rendering in Brefeld’s mostly accurate drawings, and even the slightest inflation of the sterigmata would certainly have drawn his attention in connection with the simultaneously described species of Pachysterigma J.-Olsen apud Bref. [= Tulasnella J. Schroet.]. Another notable feature of Bre-feld’s fungus seems the habitat, “kommt auf Erde, selten auf Stämmen . . . vor”, but the terrestrial growth may be typical only for the imperfect state ascribed to this species.

These considerations tend to show that the identity of T. granulata has not yet been solved, and that the problem deserves attention from European mycologists. It may belong to Botryohypochnus as defined above.

Brefeld ascribed a Botrytis-like stage to his species: he tentatively identified it with Botrytis gemella (Bonord.) Sacc. (Phymatotrichum gemellum Bonord.).

The main reason for introducing the genus Tomentella J.-Olsen apud Brefeld (1888: 9) was the existence of the conidial states ascribed to Tomentella flava and T. granulosa. Of the former species it was stated that both basidia and conidiophores were seen arising from the same hypha, and, therefore, the connection between the states has never been seriously doubted.

However, the recent description of conidial fungi that closely resemble, or may even be identical with, one of the states Brefeld described, and that have been found to belong to a discomycete, throws serious doubt upon the relations postulated by Brefeld. I intend to return to this question; for the moment it seems best not unconditionally to follow that author (or perhaps,

rather, Johan-Olsen).

8 Brefeld is reputedly unreliable as far his dimensions of microscopical structures are concerned.


BOTRYOBASIDIUM Donk restr.

Corticium sect. Botryodea Bourd. & G. in Bull. Soc. mycol. France 27: 297. 1911 (“groupe”) ; Hym. France 238. “1927” [1928]. — Type species (selected) : Corticium subcoronatum Höhn. & L. Botryobasidium Donk in Meded. Nederl. mycol. Ver. 18-20: 116. 1931.

Saprobic, neither developing into thread blights nor forming Rhizoctonia states. Fruit-body strictly

resupinate. Hyphae of wide diameter, branching rather at right angles, strongly stainable with Cotton Blue, the

ascending hyphae rather short-celled, cymosely branched, bearing the terminal basidia in more or less

candelabrum-like clusters, thus causing the granular and tufted appearance of the fruit-body when seen under

the lens. Cystidia in some species present. Basidia not much wider than supporting hyphae, hypha-like but

more or less constricted at or near middle or even urn-shaped, rather short or somewhat elongate, with

flattened apex bearing a crown of (4—) 6 (-8) tiny or small, curved sterigmata. Spores with smooth surface;

inner wall not staining orthochroma tically with Cotton Blue; not exhibiting repetition. Of several species

conidial states belonging to Oidium Link per Fr. have been established.

On rotten wood or humus.

TYPE SPECIES [selected: D. P. Rog. in Stud. nat. Hist. Univ. Iowa 17 (1) : 10. 1935].— Corticium subcoronatum

Höhn. & L. — Cf. Donk 1957: 22.

EXAMPLES.—

I. Basal hyphae not very conspicuously wider and at the same time much darker and more thick-walled than the ascending hyphae. 2. Basidia rather slender, more or less distinctly constricted at or above the middle or more definitely urn-

shaped. Subgen. Botryobasidium.—Botryobasidium subcoronatum (Höhn. & L.) Donk, B. angustisporum (Boidin) John Erikss., Pellicularia ansosa H. S. Jacks. & D. P. Rog. apud D. P. Rog.,9 Botryobasidium pilosellum John Erikss.

2. Basidia shorter, usually not distinctly constricted (spores averaging 6!). Several species known to produce conidial stages belonging to Oidium Link per Fr. Subgen. Brevibasidium John Erikss.—Botryobasidium botryosum (Bres.) John Erikss. (type of subgeneric name), B. vagum (Berk. & C. apud Berk.) D. P. Rog., B. medium John Erikss., B. conspersum John Erikss., B. obtusisporium John Erikss., Botryobasidium lembosporum (D. P. Rog.) Donk, comb. nov. (basinym, Pellicularia lembospora D. P. Rog. in Farlowia 1: 109. 1943), ? P. langloisii (Pat.) D. P. Rog.10 I. Basal hyphae much wider, darker, and thicker-walled than the ascending ones. Basidia rather distinctly constricted, almost urn-shaped. Subgen. Dimorphonema John Erikss.—Botryobasidium pruinatum (Bres.) John Erikss. (type of subgeneric name).

“Pellicularia” digitata D. P. Rog. apud G. W. Mart. (1944: 72 f. 10) has four-spored basidia that are about typically urn-shaped; it might belong to Subgenus Botryobasidium, and in that case would be the first four-spored species of the genus in the restricted sense of the present paper.

It is remarkable that of Subgenus Brevibasidium not only are most species known to produce a conidial stage but also that all these anamorphoses belong to the same form-genus, viz. Oidium Link per Fr.11, 12 :

Oidium candicans (Sacc.) Linder.—According to Linder (1942: 184), Rogers (1943: 108), and Hughes (1931: 14 f. 6) this form-species belongs to “Pellicularia” pruinata (Bres.) D. P. Rog., a

species too broadly conceived by Rogers.

9 Rogers (1943· 103 f.6) stated, sterigmata “(4-)6”. The mature basidium drawn bears 4 sterigmata, but yet looks typical of subgenus Botryobasidium.

10 Rogers (1943· 101. 3) stated in his description of this species that the basidia bear 4 sterigmata. However, his figure shows a 6-spored basidium, in shape about intermediate between this and the preceding subgenus.


The correct name for the perfect stage is, according to Eriksson (1958a: 6), Botryobasidium candicans John Erikss.

Oidium conspersum (Link per Fr.) Linder.—Linder (1942: 179) suspected that this belonged to “Peniophora fusispora”13 but Eriksson established its relationship to Botryobasidium

conspersum John Erikss. Oidium curtisii (Berk.) Linder.—Belongs to Botryobasidium vagum (Berk. & C. apud Berk.) D.

P. Rog., according to Linder (1942: 170, 203-204; “Pellicularia”14), Rogers (1943: in; “Pellicularia”), and Eriksson (1958b: 54). The latter author considerably restricted Roger’s conception of the species.

Oidium tomentosum (Berk. & C.) Linder.—Linder (1942: 205) and Rogers (1943: no) reported that this form-species had been found in one collection to be connected with Pellicularia lembospora D. P. Rog. = Botryobasidium lembosporum (D. P. Rog.) Donk.

Oidium sp.—Found connected with Botryobasidium medium John Erikss. (1958b: 54). Botryobasidium (later also called Pellicularia) started its career as a rather inclusive genus.

On the one hand it has been gradually purified by the exclusion of small entities as new generic segregates or as species transferred to other genera; on the other hand it has been constantly enriched by additions. Its limits have even been intolerably stretched, resulting in an amorphous genus indistinguishable from Corticium Fr. sensu Bourd. & G. : the inclusion for instance of the perfect stage of Sclerotium rolfsii Sacc. and of Corticium salmonicolor Berk. & Br. would appear

incongruous. The following is a list of the species of “Pellicularia” admitted to the genus by Rogers (1943) and their position suggested in the present paper.

PELLICULARIA

chordulata D. P. Rog. = Cristella sp.? isabellina (Fr.) D. P. Rog. = Botryohypochnus isabellinus (Fr. per Fr.) John Erikss. asperula D. P. Rog. = ? langloisii (Pat.) D. P. Rog. = Botryobasidium sp.? (cf. p. 26). cystidiata D. P. Rog. = Suillosporium cystidiatum (D. P. Rog.) Pouzar(1958: 31),

or, perhaps better, Jaapia sp. ochroleuca (Bres. apud W. Brinkm.) = Jaapia ochroleuca (Bres. apud W. Brinkm.)

D. P. Rog. Nannf. & John Erikss. (1933: 184) ansosa H. S. Jacks. & D. P. Rog.

apud D. P. Rog. =Botryobasidium sp.?(cf. p. 26).

11 Starting from Linder’s conception of the genus I would suggest further emendations by excluding at least the following groups: (i) Clinotrichum Cooke (name presumably not yet validly published), with Oidium lanosum (Cooke) Linder; (ii) Olpitrichum Atk., with O. macrosporum (Farl.) Linder; and (iii) Allescheriella P. Henn., with O. simile Berk, sensu Linder = Allescheriella crocea (Mont.) Hughes (1951: 1 fs. 1, 2).

12 The type species of Oidium is O. aureum (“Link” per Pers.) Link: Fr. No perfect state has as yet been found connected with it. It would be very interesting to know to which genus it belongs.

13 Linder gave as the author’s citation “Pat.” on page 170 and “(Schroet.) Hoehnel & Litsch.” on page 179. The species he had in mind would seem to be Coniophora ochroleuca Bres. apud W. Brinkm. = “Pellicularia” ochroleuca (Brinkm.) D. P. Rog. (cf. Rogers, 1943: 102). That species is certainly not a member of Botryobasidium·, it has been referred to Jaapia Bres. (see p. 28). Rogers does not mention this case at all.

14 Linder apparently erred when he stated on page 170 that O. candicans would be associable with B. vagum and O. curtisii with B. coronatum.


subcoronatum (Höhn. & L.) D.P. Rog. = Botryobasidium subcoronatum (Höhn. & L.) Donk flavescens (Bonord.) D. P. Rog. = Uthatobasidium fusisporum (J. Schroet.) Donk (cf.

p. 22). pruinata (Bres.) D.P.Rog.apudLinder

(s. str.) = Botryobasidium pruinatum (Bres.) John Erikss. lembospora D. P. Rog. = Botryobasidium lembosporum (D.P. Rog.) Donk (cf.

p. 26). vaga (Berk. & C. apud Berk.) D. P. = Botryobasidium vagum (Berk. &C. apud Berk.) D. P.

Rog. apud Linder (s. str.) Rog. koleroga Cooke sensu auct. = Koleroga noxia Donk (cf. p. 35). filamentosa (Pat.) D. P. Rog. in part. = Thanatephorus cucumeris (Frank) Donk (cf. p. 31).

Corticium praticola Kotila = Thanatephorus sp. album Dastur = Nomen dubium; apparently to be excluded.

The successive emendationsof the genus, inclusive of the one proposed above have gradually lead to a taxon that is rather sharply delimitated and recognizable not only by its general structure, of the ‘Botryobasidium’ type (and which it shares with the segregates Uthatobasidium, Thanatephorus, and Botryohypochnus), but more in particular by the shape and development of the basidia in combination with the ‘surnumerary’ number of sterigmata per basidium (averaging more than four). Botryobasidium has thus become a genus which recalls in several respects Sistotrema Fr. (cf. Donk 1956b: 4). I would not be surprised if it would appear that Botryobasidium heteronemum John Erikss. (1958a : 13) is a species of Sistotrema. What we need at the moment is a character to distinguish more easily between these two genera. This might be

found in colour tests of the hyphal walls. Some species of the Coniophoraceae have been incorporated in Botryobasidium because of

resemblance of hyphal and basidial morphology. One example is that of Coniophora ochroleuca Bres. apud W. Brinkm. which Rogers transferred to “Pellicularia”, but of which Nannfeldt & Eriksson (1953: 184) demonstrated that it should rather be included in Jaapia Bres. Another instance is Pellicularia cystidiata D. P. Rog. which essentially is nothing but another species of Jaapia, if that genus is somewhat inclusively emended. That genus then would have three species. Pouzar prefers to place each in a genus of its own, keeping Jaapia for its type, and adding Coniobotrys Pouzar (1958: 32) for Jaapia ochroleuea (Bres. apud W. Brinkm.) Nannf. & John Erikss. and Suillosporium Pouzar (1958: 31) for Pellicularia cystidiata. The difference of the Coniophoraceae (where Jaapia belongs) from Botryobasidium and the Corticiaceae in general is in the double spore wall of which the inner layer becomes strongly, orthochromatically coloured with Cotton Blue.

There are several other genera which, like Botryobasidium and Sistotrema, have basidia of more or less typical urniform shape, but these genera never show basidia with more than four

sterigmata: Galzinia Bourd., Hyphoderma Wallr. emend. Donk (1957: 13; Eriksson 1958b: 95). One species that has been referred to Hyphoderma (Peniophora polonensis Bres.) has also been

referred to “Pellicularia”, or Botryobasidium, by Boidin (1957: 121).

THANATEPHORUS Donk Corticium sect. Hypochnoidea Killerm. in Nat. PflFam., 2. Ausg., 6: 136. 1928, in part. — Type species

(selected): “C. vagum (B. et C.) Rea (= Hyp. Solani (Prill, et D.))” sensu Killerm. = Thanatephorus cucumeris (Frank) Donk.


Thanatephorus Donk in Reinwardtia 3: 376. 1956.

Mycelium often partially soil-harboured, partially parasitic and affecting herbs or herbaceous portions of

woody plants, tending to form a Rhizoctonia state consisting of fibrils or strands running over host and usually

in addition sclerotia. Fruit-body resupinate, effused, indeterminate, delicate to thin, consisting of a film or

weft of intertwined prostrate hyphae, the basidia generally formed in clusters on (poorly to) well-developed,

branched, short-celled ascending hyphae. Hyphae of fruit-body wide (the basidiferous 5-15 μ in diameter),

branching at right angles (branches slightly constricted at bases with first septa at short distances from their

bases), thin-, the basal ones often firm-, walled, colourless or the basal ones often somewhat coloured; clamps

lacking. Cystidia and gloeocystidia lacking. Basidia short and plump, barrel-shaped to obovoid, hardly or only

slightly swollen if compared with supporting hyphae, not narrowed and substipitate at bases, with rounded

tops, 2-4(-5)-spored; sterigmata comparatively stout, up to about as long as the basidia (rarely longer),

somewhat fusoidly swollen when producing the spores, not curved outwards like horns. Spores ellipsoid, ovoid,

adaxially flattened, colourless; apiculus sublateral, prominent, usually truncate; walls smooth, thin, non-

amyloid; exhibiting repetition. Conidial states lacking.

TYPE SPECIES (by original designation).—Hypochnus solani Prill. & Delacr. = Thanatephorus cucumeris (Frank)

Donk.

EXAMPLES.—Thanatephorus cucumeris (Frank) Donk (Rhizoctonia solani Kühn, Hypochnus solani Prill. &

Delacr.), see below; Hypochnus filamentosus Pat. (incompletely known, see note below) ; Corticium praticola

Kotila; Pellicularia filamentosus f. sp. timsii Exner; Corticium microsclerotia (Matz) G. W. Weber (name technically

pertaining to the rhizoctonial state) ; Hypochnus sasakii Shirai (see also note under Pachysterigma griseum

Racib., below) ; Corticium areolatum Stahel (name preoccupied) ; etc. —The specific status of most of these taxa

is not yet satisfactorily established and should come under discussion again.

This segregate from Botryobasidium Donk shares its stout sterigmata and repetition of spores with Uthatobasidium Donk and with Ceratobasidium D. P. Rog., and recalls to mind more in particular the one parasitic species of the latter genus, Ceratobasidium anceps (Bres. & H. Syd.) H. S. Jacks. The chief differences from the former genus are in the saprobic nature of Uthatobasidium and in its lack of Rhizoctonia states.

Olive (1957: 431) thinks both Uthatobasidium and Thanatephorus superfluous and, evidently relying on the sterigmata and repetition of the spores, includes the species in Ceratobasidium. Pilât (1957: 80) does the same, apparently without knowledge of the two then recently proposed genera. In so doing they oppose Martin who defends the Ceratobasidiaceae in his original circumscription: he still believes the sterigmata of Ceratobasidium on the one hand, and of ‘Pellicularia’ in part (viz. of Uthatobasidium and Thanatephorus) on the other hand, to be different organs, namely ‘epibasidia’ in the first case, and merely sterigmata in the second.

In my opinion Pilât and Olive have put too much stress on the undeniable similarity of the sterigmata and on the repetition of the spores in transferring Uthatobasidium and Thanatephorus to Ceratobasidium; they have paid too little attention to all other characters of the two genera, which very closely agree with those of Botryohypochnus as to the general structure (which is of the Botryobasidium type) and the shape of the basidia in particular, rather than of Ceratobasidium as delimited on page 20. These differentiating characters are emphasized in the key to the genera on page 17.

There are ample motives also to exclude the “Pellicularia koleroga” complex

30 FUNGUS [JAARG. 28 NO. 1-4

as currently understood as a genus of its own, Koleroga Donk (p. 35). In that taxon no soil-harboured mycelium is known to occur : it develops into typical, aerial thread blights on shrubs and trees only; the young basidia are globose and much wider than the supporting hyphae; the structure of the fruit-body is different, lacking the ascending hyphae that bear the basidia in

Thanatephorus; the sterigmata, although relatively stout, are horn-shaped (curved outwards) rather than straight and fusoidly swollen; the spores do not exhibit repetition; and so on.

In view of the actual state of our knowledge I believe that it is at least premature to treat the contents of Thanatephorus as a single species: such a thesis has not yet been proven and ignores many results of careful research that have accumulated. A cautious position is to state that with our present knowledge the delimitation of the species is still very insufficiently understood and that renewed careful taxonomic studies are much desired.

The taxonomist, especially if he is inclined to a broad species concept, will be impressed by the great morphological resemblances he encounters, and if in addition he is disposed to neglect the considerable but chaotic amount of information gathered by plant-pathologists, he will end with an all inclusive species. The usual herbarium specimens are very incomplete and should not be the sole source of information on the present genus. The solution to concentrate chaos within the limits of a single species instead of a single genus, is no real solution at all.

Another approach is not to conceal what we do not know and to point out where confusion prevails: the taxonomist should indicate where he needs information or confirmation. In any

case it appears unjust summarily to put aside conclusions that were reached after thorough and extensive research (as in the cases of Hypochnus sasakii, Corticium areolatum, and C. microsclerotia). The knowledge of the taxonomist is as yet too limited to allow a really authoritative conclusion. What he needs first is a sifting in a digestable form of the information available in the bewilderingly confusing wealth of plant-pathological literature about this genus.

Thanatephorus is an important genus among the phytopathogenic hymenomycetes, because all the species here attributed to it may cause more or less serious diseases of economically important plants. Especially plant pathologists are likely to object to the rapid changes of the generic name of these fungi—from Hypochnus, Corticium, and Rhizoctonia, etc., first to Botryobasidium, then to Pellicularia, and now again to Thanatephorus or Ceratobasidium—as highly disturbing, but from a purely taxonomic point of view such changes are unavoidable.

The ‘mycelial’ states of the species of Thanatephorus are often found without their corresponding basidiferous states and hence have been described as imperfect fungi, mostly as species of Rhizoctonia DC. per Fr., or sometimes, if stress was laid on the sclerotia, of Sclerotium

Tode per Fr., and, possibly, of Moniliopsis Ruhland. The type species of the form-genus Rhizoctonia is R. crocorum (Pers.) per Mérat : Fr., an imperfect stage of Helicobasidium purpureum Pat. = Helicobasidium brebissonii (Desm.) Donk (Auriculariaceae). Although there is much similarity between R. crocorum and the rhizoctonias of Thanatephorus it would seem defensible, and might appear profitable, to stress the


Botryobasidium-like characters of the hyphae of the mycelium and to separate the latter group from Rhizoctonia.

The form-genus Moniliopsis was published for the “Vermehrungspilz” or “maladie de la toile”, Moniliopsis aderholdii Ruhland,15 causing seed-bed or cutting bench diseases in Europe. The

genus was characterized by: “Myzel. . . in akropetaler Ordnung kettenförmige, verzweigte Monilia-ähnliche, sich nicht von einander lösende, nicht frei werdende und bald absterbende ‘Pseudokonidien’ bildend” and the presence of scierotia. The resemblance of the conidia-like cells with those of Monilia is merely superficial. These pseudoconidial chains may be initial and abortive states of the sclerotia; in any case they are structures very closely resembling such states.

Rant (1915) identified the mopo fungus causing a disease of the cinchona seed-beds in Java with Moniliopsis aderholdii. The relation of the “Vermehrungspilz” and the mopo fungus to perfect states has been sought in various directions, originally, of Botrytis cinerea Pers. per Schw.: Fr. (Sclerotinia Fuck., Discomycetes), but later, of Rhizoctonia solani (Hypochnus solani). This latter view, first clearly expressed by Duggar (1916l6), has been accepted (Simon Thomas, 1925) or rejected (Müller, 1923). If one compares the figures published by Ruhland (1908 : fs. 1, 2) of the Moniliopsis pseudoconidial chains with those of the principal elements of the Rhizoctonia solani sclerotia (Duggar, 1915: ƒ · 9 ; Peltier, 1916: fs. 3, 4), it must be admitted that the resemblance is striking indeed. Those who accept Duggar’s conclusion and also want to

assign the fungus a place in the classification of imperfect fungi will have to decide, for instance, if Moniliopsis should perhaps be merged into Rhizoctonia or the rhizoctonias of Thanatephorus taken from the latter genus and transferred to Moniliopsis.

It may be that ‘parasitic’ in the generic description will have to be altered in due time into ‘parasitic or symbiontic’, if one wants to make such distinctions, since the large majority of orchid symbionts would appear to be mycelial states of Thanatephorus. Hypochnus solani is often mentioned in this connection. For these symbionts, see especially Bernard (1909) and Burgeff (1909, 1932, 1936).

The hyphal characters of these symbionts, as well as the presence of the Monilia-like pseudoconidia and the formation of sclerotia in culture agree well with mycelium in culture of Thanatephorus cucumeris. Bernard (1909) referred them to Rhizoctonia and compared them in particular with R. solani, while Burgeff (1909: 16) called them “Orcheomycetes”, stipulating that he used this word (and its singular form “Orcheomyces”) “ohne diesem Namen eine systematische Bedeutung zuzuerkennen.” Later on he abandoned this denomination to replace it

by a still more neutral one, “Mycelium radicis” = M. R. (Burgeff, 1911). Like Bernard who entered these orchid fungi under binomial names under Rhizoctonia, Burgeff (1932) finally also resorted to binomials under Rhizoctonia for those species resembling Rhizoctonia solani.

THANATEPHORUS CUCUMERIS.—This species is generally known as Rhizoctonia

15 The correct name for the imperfect state only should apparently be derived from Mor tierella arachnoides Therry & Thierry (1882: 161), “Araignées des serres”.

16 See this publication for previous literature on the “Vermehrungspilz”.


solani Kühn if stress is laid on the sterile condition, or as Hypochnus solani Prill. & Delacr. if the basidiferous state is emphasized. For some time the denomination Corticium vagum Berk. & C. was often used but this was erroneous because the type of that name proves it to belong to a species of Botryobasidium, viz. B. vagum (Berk. & C. apud Berk.) D. P. Rog. The use now en

vogue of the name Pellicularia filamentosa (Pat.) D. P. Rog. is in my opinion not defensible, first, because its conspecificity with Hypochnus solani is still questionable (see below) ; secondly, because the name Pellicularia Cooke is impriorable and thus not available for a genus of Hymenomycetes : compare Donk (1954); and, thirdly, there is an earlier name (Hypochnus cucumeris Frank) available.

The identity of Hypochnus solani with the earlier described Hypochnus cucumeris Frank (1883a) is, in my opinion, quite obvious. Frank mentioned also many microscopical details (1883b: 524). This conclusion was already previously reached by Nakata (1926: 14, 19) and by Drs J. A. Nannfeldt and J. Eriksson (oral communication). As to the symptoms Frank described, these are clearly the same as those caused by typical Hypochnus solani. The following passages from his original account give a good picture of the fungus itself:—

“ ... dem unbewaffneten Auge sehr auffallender Pilz ... welcher die untersten Theile der Pflanze überzieht. Von der Oberfläche des Bodens, also auf dem dort liegenden oberen Wurzelende beginnend, bedeckt meist mehrere Centimeter weit am Stengel und wohl auch an den untersten Blattstielen emporsteigend eine faserige graue oder braünlichgraue Pilzhaut die genannte Organe, mit ihrem flockig oder strahlig aufgelösten oberen Rande allmählig auf den Theilen weiter aufwärts kriechend. Diese Haut ist das Mycelium des Pilzes, welches sich in seinen älteren Theilen auf der ganzen Aussenfläche mit einem Hymenium bedeckt, indem überall von der Hyphen aus mehr oder weniger dicht beisammenstehende längliche Basidien sich abzweigen, welche nach Hymenomycetenart auf ihrer Spitze an vier Sterigmen je eine einzellige, ovale, farblose Spore abschnüren. Duch die Sporen erscheint die Oberfläche des Hymeniums pulverig ... Das Mycelium liegt den genannten Theilen der Gurkenpflanze ohne jede Verbindung lose auf, lässt sich leicht davon abziehen, und die Theile erscheinen darunter im allgemeinen frisch und gesund. Aber an einer Stelle der untersten Theile findet man den Pilz in das Innere eingedrungen ...”—Frank (1883a: 63).

Frank also supplied the following microscopical description of the fruit-bodies producing

basidiospores.

“...Die Myceliumhaut besteht aus sehr locker verflochtenen Hyphen, welche 0,006 bis 0,009 mm Durchmesser, zahlreiche Querscheidewände haben, sich seitlich verzweigen und einen mehr oder minder geschlängelten in verschiedenen Richtungen gehenden Verlauf haben; ihre Membranen sind ziemlich dick, farblos oder blassbraun ... Die Entwickelung des Hymeniums auf dem Mycelium besteht darin, dass an den oberflächlich verlaufenden Mycel-hyphen, die dann meist reichlich Querscheidewände bekommen, so dass ihre Glieder ziemlich kurz, nur wenig länger als breit sind, von vielen dieser Gliederzellen an der nach aussen ge-kehrten Seite kurze seitliche Aestchen getrieben werden, welche meist wiederum eine oder einige Querscheidewände bekommen und büschelig sich in kurze Zweige verästeln. Die ovalen äussersten Zweigzellen stellen die Basidien dar; sie sind meist in ziemlich grosser Anzahl dicht beisammenstehend mit ihren Spitzen nach aussen gerichtet. Auf diesen Spitzen treiben sie nach Hymenomyzeten-Art vier dünne Sterigmen, auf denen je eine Spore abgeschnürt wird. Die Sporen sind oval, durchschnittlich 0,009 mm im längeren Durchmesser, einzellig, farblos. Sie keimen auf Wassertropfen schon nach 24 Stunden, indem sie aus einem oder beiden Enden einen Keimschlauch treiben. ...”—Frank (1883b: 524).

It is easy to understand that one is reluctant to take up still another name for Hypochnus solani; it was even to be expected that it would be rejected without much argument (Olive 1957: 431), but yet, in my opinion, it will be


difficult—if not impossible—to suggest a reasonable alternative to the identification of Hypochnus cucumeris with H. solani. Several mycologists whom I have questioned about this matter, and I myself, do not hesitate in the least confidently to accept that identification.

Thanatephorus cucumeris is currently interpreted as a highly polymorphous species

consisting of numerous, often rather different strains. Yet, in the symptoms of the diseases it causes on its innumerable hosts and in its behaviour it seems consistent enough, at least in Europe. It may be called a ‘subterranean’ rather than an ‘aerial’ fungus. It infests and parasitizes the underground parts (Rhizoctonia state) and at most the lower portion of the stalk adjacent to the soil; on these latter parts it often develops its perfect state (“collar fungus”). These fruit-bodies spread out only to a limited extend from the root-crown upwards and laterally on under-surfaces of the host. Sometimes fruits and the like touching the infected soil become the substratum of the basidiferous state, always only at the contacting side. Exner (1953: 704) reported that only 'Corticium solani’ was isolated in nature from roots or lesions on basal stems, which might indicate that the three other taxa of Thanatephorus reported from Louisiana, U.S.A., were confined to aerial parts of hosts attacked.

The plants attacked are herbs, or woody plants still more or less in a herbaceous stage (seedlings), or more rarely herbaceous parts of woody plants. The symptoms of the diseases caused are manifold and may vary from host to host. The parts directly affected are primarily subterranean or such as are close to the soil; if these are diseased secondary symptoms may be

shown by aerial parts and may be usually explained as caused by malnutrition because the roots or the stem have been seriously damaged. The many diseases are well known to the phytopathologist and can be found dealt with in every textbook on plant diseases. Some of the best known types are damping off, or belong to the classes of stem rots (parts near or in the soil), potato scab (in various forms), root rots (inclusive of fleshy roots), fruit rots (egg plant: Wolf, 1914)· Leaf rots are also known on suitable hosts, with the rosette habit. For an important summary on earlier literature see, for instance, Duggar (1915) and for a detailed account, Peltier (1916).

Rhizoctonia solani Kühn, the mycelial state, was finally brought into connection with the correct basidiferous state, viz. Hypochnus solani, which species was at first erroneously referred by Burt (Rolfs, 1903) to Corticium vagum, a saprobic species belonging to Botryobasidium sensu stricto. For important historical reviews on the attempts to connect the Rhizoctonia state with its perfect condition, see Duggar (1915) and Braun (1930)·

The Rhizoctonia state produces its sclerotia in various degrees of profusion. The potato tuber

is one of the seats on which they are generally copiously formed. On many hosts, however, sclerotial formation is relatively rare. In cultures they are invariably formed under suitable conditions. They may vary considerably in size, shape, and other characters.

As to their structure, the sclerotia are hardly typical, if such bodies are characterized as having an outer, distinctly differentiated rind. The latter is lacking. For accounts of their internal structure, see, for instance, Duggar (1915: 439-443 fs- 7-9)·


HYPOCHNUS FILAMENTOSUS Pat. apud Pat. & Lagerh. in Bull. Soc. mycol. France 7: 163 pl. 11 f. 2. Sept. 30, 1891. — Pellicularia filamentosa (Pat. apud Pat. & Lagerh.) D. P. Rog. in Farlowia 1: 113 f. 11. 1943, in part.

Rogers (1943: 113-115), who studied Patouillard’s type, concluded that Hypochnus

filamentosus was identical with H. solani and that the “similarity of H. filamentosus and H. solani would seem sufficiently apparent from the illustrations, published in successive numbers of the same journal; it is a wonder that Prillieux & Delacroix’s account of H. solani was ever allowed to appear, and a greater wonder that their name was not at once reduced to synonymy.”

One thing about Hypochnus solani Prill. & Delacr. we know, and should not reject as unimportant without sound arguments, is that it is a soil fungus. This knowledge we lack altogether in connection with H. filamentosus. That species is that rare thing in Thanatephorus of which we know only the basidiferous fruit-body—and nothing else. It was described from Ecuador from living leaves of Dianthus caryophyllus and Amaryllis; one fruit-body half-way up a Dianthus leaf was depicted; no symptoms were indicated. Drs J. A. Nannfeldt and J. Eriksson drew my attention to the word ‘pink’ used in describing the fruitbody, “roseus vel albidus”, a colour never recorded for H. solani. I do agree with Rogers that the structure of the fruit-body as depicted by Patouillard agrees well with that of H. solani (= Th. cucumeris), but in Thana-tephorus this is hardly enough. I prefer to list it as an as yet insufficiently known species and, therefore, the name H. filamentosus as a nomen dubium.

PACHYSTERIGMA GRISEUM Racib., Paras. Algen Pilze Java’s 1: 30. 1900.— Tulasnella grisea (Racib.) Sacc. & P. Syd. in Sacc., Syll. Fung. 16: 203. 1902.

Described from the leaves of Eichhornia crassipes at Buitenzorg (= Bogor), Java, causing necrotic spots. The description, which is rather extensive, gives an excellent picture of a species of Thanatephorus. Rogers (1933: 202), who studied the Bogor specimen, concluded that the hyphae are those of the parasitic species of Corticium (= Thanatephorus), but he could not find basidia and spores. There is no doubt that Raciborski was struck by the stout sterigmata and,

therefore, placed his species in Pachysterigma Bref. (= Tulasnella J. Schroet.), although he described them merely as “ziemlich lang (bis 5[!] μ).” It also is quite likely that the other fungus he mentioned from the spots on the host is nothing but the Rhizoctonia state of Pachysterigma griseum, “bald von [Pachysterigma griseum) getrennt, bald durcheinander wachsend zu finden, und zwar ein runde, braune, bis 1 mm. breite Sclerotien bildender, den ich nicht bestimmen konnte”.

The descriptions of the lesions caused reminds one strongly of those described for Hypochnus sasakii Shirai by Matsumoto & Hirane (1933) and obtained on leaves of Eichhornia crassipes after artificial infection. It may be that Pachysterigma griseum is the first strain of Hypochnus sasakii described under a validly published name for the perfect state, but there is one item that prevents the assumption of conspecificity without additional knowledge of Raciborski’s fungus : he described the spores as ovoid, 5 x 4 μ, which is far too small for

Hypochnus sasakii. I have not been able to collect additional specimens at Bogor.


Koleroga Donk, gen. nov.17

Misapplication.—Pellicularia Cooke sensu Höhn, in S.B. Akad. Wiss. Wien (Math.-nat. Kl., Abt. I) 119: 395. 1910 (transferred to Corticium) ; D. P. Rog. in Farlowia 1: 96. 1943, in part.

Parasitica, mycelio filiformi; fructificationes paginam inferiorem laminarum arborum vivarum obtegentes, e hyphis laxe intertextis repentibus formatae, hyphis ipsis ad ramificationes brevissimas vel breves basidia solitaria vel pauca congeste gerentibus, 5-10 μ diam., rectangulo-ramificatis, efibulatis. Cystidia desunt. Basidia immatura subglobosa, hypha sustinentia manifeste latiora, matura sterigmata 2-4 satis longa et curvata formantia. Sporae oblongae, unilateraliter subdepressae, incolores, laeves, nunquam denuo sporas repetentes.

Thread blights on woody plants (trees and shrubs). Spreading from stem and branches over the under-

surfaces of leaves and there forming a film or weft which may become basidiferous: killed leaves of hosts

breaking off and dangling for some time, being suspended by mycelial threads. Fruit-body consisting of

intertwined prostrate hyphae which produce the basidia often solitary or in twos on very short, or in poorly

developed clusters on branched, not typically ascending side-branches. Hyphae of fruit-bodies rather wide

(about 5-10 μ in diameter), branching at right angles, thin- to firm-walled, colourless or the basal ones often

becoming slightly coloured; clamps lacking. Cystidia and gloeocystidia lacking. Basidia nearly globular when

young and distinctly wider (2-3 x) than supporting branches, short and plump and short-barrelshaped when

mature, 2-4(-6)-spored18; sterigmata comparatively long, but shorter than basidial body, slender-conical and

horn-shaped (curved outwards). Spores ovoid or oblong to somewhat fusiform or subcylindrical and rather long

(7-16 μ), adaxially slightly to distinctly flattened or even somewhat depressed, colourless ; walls smooth, non-

amyloid, slightly but distinctly thickened and refractive (according to Rogers) ; not exhibiting repetition.

Conidial states lacking.

Tropical and subtropical; developing under hot and humid conditions.

TYPE SPECIES.—Koleroga noxia Donk. Genotype specimen: that portion of the type of Pellicularia Koleroga

Cooke (K) represented by the hyphae giving rise to basidia and spores illustrated by Burt (in Ann. Missouri bot.

Gdn 5: 124 f. 1a. 1918; 13: 293 f. 1a. 1926).

EXAMPLES.—

I . Sclerotia present on the rhizomorphic threads, conspicuous, although sometimes rare. America ............................................................................................................................Corticium stevensii Burt

I. Sclerotia of this type absent; films on under-surfaces of leaves may have a finely mottled appearance due to minute hyphal conglomerations. Asia and America. Koleroga noxia19

The differences of Koleroga from Thanatephorus have already been discussed in connection with the latter genus (p. 30).

The genus is in need of a critical survey of both the taxonomic and the phytopathological literature. In contrast to Rogers (1943: 112) I am not at all convinced that only a single species is

involved and for the time being would rather admit two species as differentiated above. The establishment of the correct names for these species proved to be an exceptionally knotty nomen- clatural problem, the solution of which depends to some degree on the range of one’s conception of the species. A more careful analysis of the species is postponed until I have gathered more information about them.

17 Kole roga, an Indian name, used in Mysore, meaning black rot. Gender: f.

18 Six-spored basidia (“rarely”) were observed by Wolf & Bach (1927: 701).

19 Koleroga noxia Donk, sp. nov. Sclerotia nulla. Fructificationes plerumque glomerulis hypharum minutis variegatae. — Type: same as of generic name.


Two other species should be taken into consideration, Corticium invisum Petch (1925: 316), spores “5-6 x 3-4 µ”, on various hosts, i.a. tea, “on which it causes the disease known as Black Rot in Ceylon;” and C. pervagum Petch (l.c.) on Erythroxylon cola L. Specimens referred to the latter species by Petch at Kew may appear to be close to Koleroga noxia.

REFERENCES

The following titles have been cited by their dates printed in italics.

BERNARD, N. (1909) in Ann. Sci. nat. (Bot.) IX 9: 1-196. BOURDOT, H. & A. GALZIN (1928), Hym. France. “1927”. BRAUN, H. (1930), Wurzeltöter Kartoffel. (Monogr. PflSchutz 5). BREFELD, O. (1888), Unters. Gesammtgeb. Mykol. 8. “1889”. BRESADOLA, G. (1911) in Ann. mycol. 9: 425. BURGEFF, H. (1909), Wurzelpilze Orchid.; (1911), Anzucht trop. Orchid. Samen; (1932), Saprophyt. u. Symb.;

(1936) Samenkeimung Orchid. DONK, M. A. (1931) in Meded. Nederl. mycol. Ver. 18-20; (1954) in Reinwardtia 2: 425-434; (1956a) in

Reinwardtia 3: 363-379; (1956b) in Fungus 26; (1957) in Taxon 6; (1958) in Blumea, Suppl. 4. DUGGAR, B. M. (1915) in Ann. Missouri bot. Gdn 2: 403-458; (1916) in Ann. Missouri bot. Gdn 3: 1-10. ERIKSSON, JOHN (1958a) in Svensk bot. Tidskr. 52; (1958b) in Symb. bot. upsal. 16 (1); (1958c) Stud. Swedish

Heterob. Aphyll. EXNER, B. (1953) in Mycologia 45. FRANK, B. (1883a) in Ber. dtsch. bot. Ges. 1: 62-63; (1883b) in LandwSch. Jb. 2. HÖHNEL, F. X. R. VON & V. LITSCHAUER (1906) in S.B. Akad. Wien (Math.-nat. Kl., Abt I) 115; (1907) in S.B. Akad.

Wien (Math.-nat. Kl., Abt. I) 116; (1908) in S.B. Akad. Wien (Math.-nat. Kl., Abt. I) 117. HUGHES, S. J. (1951), Stud, micro-fungi. VII. (Mycol. Pap., C.M.I. No. 41). JACKSON, H. S. (1949) in Canad. J. Res. G 27; (1950) in Canad. J. Res. C 28. LINDER, D. H. (1942) in Lloydia5. MARTIN, G. W. (1939) in Mycologia 31; (1941) in Lloydia 4; (1948) in Lloydia 11; (1957) in Brittonia 9. MATSUMOTO, T. & S. HIRANE (1933) in J. Soc. trop. Agr., Formosa 5: 367-373. MÜLLER, Κ. Ο. (1923) Arb. biol. Reichsanst. Land- u. Forstwsch. 11: 321-325. NAKATA, K. (1926) in Bull. sci. Fak. Tercult. Kjusu Univ. 2 (1). NANNFELDT, J. A. & J. ERIKSSON (1953) in Svensk bot. Tidskr. 47. OLIVE, L. S. (1957) in Amer. J. Bot. 44. PARKER-RHODES, A. F. (1954) in Trans. Brit. mycol. Soc. 37. PELTIER, G. L. (1916) in Bull. Illinois Exp. Stat. No. 189: 279-390. PETCH, T. (1925) in Ann. roy. bot. Gdns, Peradeniya 9. PILAT, A. (1957) in Česka Mykol. 11. POUZAR, Z. (1958) in Česka Mykol. 12. RANT, A. (1915) in Bull. Jard. bot. Buitenzorg II No. 18. ROGERS, D. P. (1933) in Ann. mycol. 31; (1935) in Stud. nat. Hist. Univ. Iowa 17 (1); (1943) in Farlowia 1: 95-

118. ROGERS, D. P. & H. S. JACKSON (1943) in Farlowia 1. ROLFS, F. M. (1903) in Science II 18: 729. RUHLAND, W. (1908) in Arb. biol. Anst. Land- u. ForstwSch. 6: 71-76. SIMON THOMAS, K. (1925), Onderz. Rhizoctonia. THERRY, J. J. & THIERRY (1882) in Rev. mycol. 4. WAKEFIELD, E. M. (1952) in Trans. Brit. mycol. Soc. 35. WOLF, F. A. (1914) in Mycol. Gbl. 4: 282-286. WOLF, F. A. & W. J. BACH (1927) in Phytopathology 17.

TAXONOMICAL NOTES ON MOLLISIACEOUS FUNGI—VI

The genus Pyrenopeziza Fuck.

J. GREMMEN

Forest Research Station, “De Dorschkamp”, Wageningen

A historical review of the genus Pyrenopeziza Fuck, is given. It appears that Pyrenopeziza takes precedence over Mollisia (Fr.) Karst. The diagnosis of the genus is emended. A subdivision based on differences of the receptacle is attempted, and eight new sections and two subsections are proposed.

Introduction.—In the course of the study of the mollisiaceous fungi it soon appeared that asci and ascospores are of little use for the identification of the species. Measuring of the spores is helpful only in a limited number of long-spored species. The receptacle, however, shows characteristic features which may be of taxonomic value, but the variation of its elements should be interpreted very carefully, since young and old receptacles may be totally different within the same species. In Pyrenopeziza rubi (Fr.) Rehm for instance, when growing on its natural substratum of raspberry stems, young excipular cells may be seen to range from 4-8 μ in diameter, whereas fully developed excipular cells measure 8-15 μ (Gremmen, 1954). In Pyrenopeziza artemisiae (Lasch) Rehm the variability of the excipular cells may even be studied

in herbarium material (Gremmen, 1955). It should also be born in mind that the excipular cells of apothecia grown in culture may differ from those of apothecia found in nature. Apothecia of Pyrenopeziza plantaginis Fuck, grown in vitro have excipular cells larger than those of apothecia developed on their natural medium. Similarly, ascospores produced in vitro are longer (Gremmen, 1952). However, in spite of certain limitations, the receptacle will be used largely as a basis for the following division of the genus Pyrenopeziza.

For the study of the present genus to be successful, it is first of all necessary to investigate such species as are known to form abundant apothecia in nature, since that alleviates the comparative study of the various developmental stages of the exciple. Species of which few apothecia can be found, should, for the time being, be avoided. Such species which Nannfeldt

(1932) called “verirrt” in general defy any attempt at identification.

The following abbreviations denote institutions and private collections where material mentioned in this

paper is located. BM: British Museum, Natural History, London; C: Botanical Museum, Copenhagen; G:

Conservatoire et Jardin Botaniques, Genève; Grd: herbarium W. D. Graddon, Congleton; Grm : author’s

herbarium; H: Botanical Museum, Helsinki; K: The Herbarium, Royal Botanic Gardens, Kew; L: Rijks-

herbarium, Leiden ; PC: Muséum National d’Histoire Naturelle, Laboratoire de Cryptogamie, Paris; PR:

Botanical Department of the National Museum, Prague; S: Naturhistoriska Riksmuseum, Stockholm; UPS:

Universitetets Institution for Systematisk Botanik, Uppsala, and ZT : Institut für Spezielle Botanik der

Eidgenössischen Technischen Hochschule, Zürich.

Historical review.—Pyrenopeziza was first used by Fuckel (1869:293), whereas the name

Mollisia originated from Elias Fries (1822:116), who by it


indicated a tribus of his genus Peziza. This tribus was raised to generic rank by P. A. Karsten (1871:15), and two sections were distinguished, viz. “Sect. I. Apothecia subiculo nullo vel minus distincto insidentia” and “Sect. II. Apothecia subiculo distincto subtomentoso insidentia Tapesia Fr. p.pr.”

Phillips (1887) regarded Mollisia sensu Fr. as a good genus, dividing it in the subgenera Niptera Fuck., Pyrenopeziza Fuck., Dilutella Phill., Mollisiella Phill., Hysteropeziza Rabh., Pseudopeziza Fuck., and Peristomialis (without author’s name). In part this scheme is based on the apothecial structure, e.g. Niptera being characterized by “texture rather firm; cups plane” and Pyrenopeziza by “texture soft; cups globose (mostly black),” but on the whole the genus is very heterogeneous.

Rehm (1896) recognized in his Mollisieae the group Eumollisieae with “Apothecien von Anfang an sitzend” and the Pyrenopezizeae characterized by “Apothecien zuerst in die oberen Zellschichten eingesenkt, durch dieselben hervorbrechend und dann sitzend.” The Eumollisieae are split up into (a) genera without a subiculum, comprising Mollisia Fr., Niptera Fr., Belonidium Mont. & Dur., and Belonopsis Sacc., and (b) genera with a subiculum such as Tapesia Pers. and Trichobelonium Sacc. The Pyrenopezizeae comprise the Pseudopezizeae with the genera Pseudopeziza Fuck., and Fabraea Sacc., and the Eupyrenopezizeae with Pyrenopeziza Fuck., Pirottaea Sacc. & Speg., Beloniella Sacc., and Velutaria Fuck.

Boudier (1907) mentioned in his family Mollisiacées the following genera: Pyrenopeziza

Fuck., Ephelina Sacc., Pirottaea Sacc., Coronellaria Karst., Mollisia Fr., Tapesia Pers., Niptera Fr., Mollisiella Boud., and Spilopodia Boud. He characterized Mollisia as follows: “L’aspect général aplati, plus étalé, des réceptacles qui sont toujours sessiles, . . . ” and Pyrenopeziza: “Les réceptacles sont généralement noirs extérieurement avec la marge plus blanche.” According to Boudier the paraphyses in Mollisia are filled with an oily protoplasm, lacking in Pyrenopeziza.

In 1912 Rehm revised the Pyrenopezizeae and the genus Pyrenopeziza was again discussed in detail.

Nannfeldt (1932) recognized two subfamilies in his Mollisioideae, viz. the Mollisieae and the Pyrenopezizeae. The first category dealt with the species having the “Apothecien ganz oberflächlich oder nur mit zapfen-förmiger Basis im Substrat eingesenkt,” those of the second group having the “Apothecien im Substrat eingesenkt, hervorbrechend.” In the former are found the genera Haglundia Nannf., Belonium Sacc., Coronellaria Karst., Tapesia Fuck., Tricho-belonium Rehm, Mollisia (Fr.) Karst., and Belonopsis Rehm; in the latter Pirottaea Fuck, and Hysteropezizella v. Höhn.

The present author already demonstrated that it is hardly possible to delimit the genus Pyrenopeziza Fuck, from Mollisia (Fr.) Karst, merely on the basis of its erumpent behaviour during development. Nannfeldt occasionally arrived at the same conclusion (1932: 123,156), but probably maintained the separation for practical purposes. Gremmen (1954) united both genera, assuming Mollisia Fr. to be the correct name. It appears, however, that Pyrenopeziza Fuck, was published two years prior to Mollisia (Fr.) Karst., so the former takes precedence. Some corrections and additions to the diagnosis previously published appear necessary.

Nov. 1958] J. Gremmen: On mollisiaceous Fungi—VI 39

PYRENOPEZIZA Fuck. em. Gremmen

Pyrenopeziza Fuck., Symb. myc. 293. 1869. Peziza ser. Ill Phialea Trib. XI Mollisia Fr., Syst. myc. 2:116. 1822. — Mollisia (Fr.) Karst., Myc. fenn. 1:15.

1871. -— Mollisia Fr. sensu Gremmen in Fungus 24:2. 1954.

Apothecia scattered or gregarious, small, ceraceous, sessile, shortly stipitate or broadly based, erumpent or

superficial from the beginning, cupuliform to disciform. Receptacle even, verrucose, fimbriate or hairy. Ectal

excipulum consisting of textura globulosa, i.e. of roundish, oblong or polyhedral cells which may be variously

coloured or colourless; marginally with or without elongate or club-shaped, coloured or colourless cells.

Hymenium variously coloured. Hypothecium colourless, undefined. Asci minute, clavate or cylindrical.

Ascospores 4 or 8 per ascus, 1-celled, or 2- or 4-septate, bacilliform, cylindrical, ovoid, comma-shaped or

acicular. Paraphyses filiform or lanceolate, colourless. Type species: Peziza chailletii Pers. (vide Nannfeldt, 1932:

137).

Already Nannfeldt distinguished certain groups in the genus Pyrenopeziza, one of which is “ ... u.a. charakterisiert durch verhältnismässig grosse, mehr oder weniger niedergedrückt ellipsoidische Apothecien,...” A number of species were considered to belong to this group. Another group” ... bildet eine Anzahl von Arten, die sämtlich auf vorjährigen Stengeln wachsen und deren Apothecien mit breiter Basis ihren Holzzylindern aufsitzen.”

As appears from the following it is possible to distinguish eight sections which are mainly based on differences of the colour and structure of the receptacle.

KEY TO THE SECTIONS

I a. Paraphyses filiform 2a. Receptacle even, glabrous

3a. Receptacle dark brown or red-brown 4a. Receptacle sessile on a hypostroma . . Sectio 2. Hypostromatina 4b. Receptacle without a hypostroma . . Sectio 3. Astromatina

3b. Receptacle differently coloured 5a. Receptacle olivaceous or greenish brown…………….. Sectio 4. Elaiodiscina 5b. Receptacle melleous or light brown . ………………….. Sectio 5. Melipeziza

2b. Receptacle verrucose, fimbriate or hairy 6a. Receptacle verrucose

7a. Ascospores bacilliform Sectio 1. Pyrenopeziza, subsection a. 7b. Ascospores ovoid .................................................. Sectio 1. Pyrenopeziza, subsection b.

6b. Receptacle with appendages 8a. Receptacle fimbriate ........................................ …..Sectio 6. Fimbriaria 8b. Receptacle hairy ................................................... Sectio 7. Trichantina

I b. Paraphyses lanceolate Sectio 8. Hemipirottaea

Sectio I. PYRENOPEZIZA

Apothecia erumpentia, sessilia vel basi stipite fusca. Receptaculum verrucosum. In caulibus anni praeteriti emortuis. Sectionis species typica: P. chailletii (Pers.) Fuck.

Apothecia erumpent, with or without dark brown base submerged in host. Receptacle verrucose, owing to

presence of groups of protruding cells. Inhabiting last year’s stems. Type species: P. chailletii (Pers.) Fuck.

The section is divided in two subsections.


Subsectio a. Bacillares Gremmen, subsect. nov.

Ascosporae bacilliformes, unicellulares vel pluricellulares. Ascospores bacilliform, 1 - or more-celled.

1. PYRENOPEZIZA CHAILLETII (Pers.) Fuck., Symb. myc. 294. 1869.

Peziza chailletii Pers., Myc. eur. 1: 288. 1822 (basionym).

Material: Pyrenopeziza chailletii Fuck. (G). Host : Chaerophyllum aureum.

2. PYRENOPEZIZA POTENTILLAE (Rostr.) Nannf. in Svensk bot. Tidskr. 22 : 136. 1928.

Trochilapotentillae Rostr. in Medd. om Grönl. 3: 540. 1888 (basionym).

Material: Trochila potentillae Rostr., type (C). Host: Potentilla nivea.

3. PYRENOPEZIZA THALICTRI (Peck) Sacc., Syll. Disc. 360. 1889.

Peziza thalklri Peck in Ann. Rep. N.Y. State Mus. 29: 55. 1878 (basionym). — Pyrenopeziza osiliensis Vgr. in

Bot. Not. 40. 1900. — Beloniella osiliensis (Vgr.) Rehm in Ber. bayr. bot. Ges. 13: 181.1912.

Material: no. 517, Pyrenopeziza thalictri (Grd). Host: Thalictrum spec.

4. PYRENOPEZIZA PYRENOCARPOIDES Rehm in Ber. bayr. bot. Ges. 13: 175. 1912.

Mollisia pyrenocarpoides (Rehm) Gremmen in Fungus 25: 9. 1955.

Material : Pyrenopeziza pyrenocarpoides Rehm (S). Host: Cirsium spec.

5. Pyrenopeziza buniadis (Nannf.) ex Gremmen, comb. nov.

Mollisia buniadis Nannf. spec. nov. inedit, (basionym).

Material: Flora suec. 1913, Mollisia buniadis Nannf., Uppsala, Slottsbacken, ii. 6. 1925, leg. J.

A. Nannfeldt, type (UPS).

Host: Bunias orientalis.

Apothecia 400—500 μ diam., basi 80 μ lata. Receptaculum verrucosum, melleum, e textura globulosa.

Cellulae excipuli 7-10 μ diam. Margo cellulis minutis formata. Asci 38-50 x 5.5 µ. Ascosporae 9.5-10.5 x 2 µ,

unicellulares, incoloratae. Paraphyses filiformes, incoloratae, guttulatae.

Apothecia 400-500 μ across, with 80 μ broad base. Receptacle consisting of textura globulosa, honey-

brown, verrucose owing to presence of protruding cells. Excipular cells 7-10 μ in diameter, marginally with

short cell-excrescences. Asci 38-50 x 5.5 μ. Ascospores 9.5-10.5 x 2 μ, colourless, 1-celled. Paraphyses

colourless, filiform, filled with numerous, minute guttulae.

6. PYRENOPEZIZA LYCHNIDIS (Sacc.) Rehm in Krypt. Fl. 3: 1265. 1896.

Pyrenopeziza sphaerioides (Pers.) Fuck. f. lychnidis Sacc., Syll. Disc. 365. 1889.

Material: Rabenh.-Pazschke, Fgi. eur. & extraeur. 4173, Pyrenopeziza lychnidis Sacc. (L).

Host: Lychnis spec.

7. PYRENOPEZIZA LAVATERAE E. Müller in Sydowia 9: 239. 1955. Material: Fgi. Pakistan 4387,

Pyrenopeziza lavaterae E. Müll., type (ZT). Host : Lavatera spec.

Nov. 1958] J. Gremmen: On mollisiaceous fungi—VI 41

Subsectio b. Ovoideae Gremmen, subsect. nov.

Ascosporae ovoideae vel formam commatis aemulantes, unicellulares.

Ascospores ovoid or comma-shaped, 1-celled.

8. Pyrenopeziza nigrella Fuck., Symb. myc. Nachtr. 3: 30. 1871.

Material: Pyrenopeziza nigrella Fuck. (G). Host: Succisa pratensis. 9. Pyrenopeziza lythri Fautr. in Rev. myc. 14: 3. 1892. Material: Rabenh.-Pazschke, Fgi. eur. & extraeur. 4470, Pyrenopeziza lycopodis f. lythri Rehm (L). Host: Lythrum salicaria. 10. Pyrenopeziza lini (Rostr.) Petr. & Syd. in Ann. myc. 22: 356. 1924.

Laestadia lini Rostr. in Bot. Tidsskr. 28: 217. 1907 (basionym).

Material: Pyrenopeziza lini (Rostr.) Petr. & Syd. (ZT). Host: Linum viscosum. 11. Pyrenopeziza bubâkii Klika in Ann. myc. 24: 136. 1926. Material: Krypt. exs. 955, Mollisia atrocinerea sensu Nannfeldt (L). Host: Dipsacus spec. Sectio 2. Hypostromatina

Gremmen, sect. nov. Apothecia ex hypostromate fusco distincto exeuntia, ceterum iis sectionis Astromatinae simillima. In

ramulis emortuis lignoque putrido. Sectionis species typica: P. caespiticia (Karst.) Gremmen.

Apothecia formed on well-developed dark brown hypostroma, in other respects resembling those of

section Astromatina. Inhabiting dead branches or decayed wood. Type species: P. caespiticia (Karst.) Gremmen.

12. Pyrenopeziza caespiticia (Karst.) Gremmen, comb. nov. Peziza caespiticia Karst., Mon. Pez. 159. 1869 (basionym). — Mollisia caespiticia (Karst.) Karst., Myc. fenn. 1:

188. 1871.

Material: Peziza caespiticia Karst., type (H). Host: Sambucus. 13. Pyrenopeziza caesia (Fuck.) Gremmen, comb. nov.

Niptera caesia Fuck, in Jahrb. Nass. Ver. Nat. 25/26: 335. 1871 (basionym). — Mollisia caesia (Fuck.) Sacc.,

Syll. Disc. 340. 1889.

Material: Niptera caesia Fuck., type (G). Host: Fagus, Salix, Tilia. Sectio 3. Astromatina

Gremmen, sect. nov.

Receptaculum fuscum vel ferrugineum, cellulis globosis vel piriformibus vel papillatis. Sectioni

Hypostromatinae simillima, sed hypostromate nullo. In caulibus anni praeteriti ramulisque emortuis. Sectionis

species typica: P. revincta (Karst.) Gremmen.

Receptacle dark brown or red-brown, excipular cells globular, pear-shaped or papillate. Resembling

section Hypostromatina, but hypostroma lacking. Inhabiting last year’s stems, and decaying branches. Type

species: P. revincta (Karst.) Gremmen.


14. Pyrenopeziza revincta (Karst.) Gremmen, comb. nov.

Peziza revincta Karst., Mon. Pez. 157. 1869 (basionym). — Mollisia revincta (Karst.) Rehm in Krypt. Fl. 3:

1264. 1896.

Material: Peziza revincta Karst., type (H). Host: Rubus, Ulmaria, Epilobium, and various Compositae.

15. Pyrenopeziza benesuada (Tul.) Gremmen, comb. nov.

Peziza benesuada Tul. in Bot. Zeit. 11: 55. 1853 (basionym). — Mollisia benesuada (Tul.) Phill., Man. Brit.

Disc. 174. 1887.

Material: Flora suec. 9452, Mollisia cf. benesuada (UPS). Host: Alnus spec.

16. Pyrenopeziza adenostylidis (Rehm) Gremmen, comb. nov. Mollisia adenostylidis Rehm in Krypt. Fl. 3: 526, 1896 (basionym).

Material: Rehm, Ascom. 49, Mollisia adenostylidis Rehm (S). Host: Adenostyles spec.

17. Pyrenopeziza arundinacea (DC.) Gremmen, comb. nov.

Xyloma arundinacea DC., Fl. France 6: 162. 1815 (basionym). — Mollisia arundinacea (DC.) Phill., Man. Brit.

Disc. 177. 1887.

Material: Rabenh. 423, Peziza cinerea Batsch (L). Host: Phragmites communis. Sectio 4. Elaiodiscina Gremmen, sect. nov.

Receptaculum olivaceum vel virido-fuscum. In caulibus anni praeteriti emortuis. Sectionis species typica:

P. carduorum Rehm.

Receptacle olivaceous or brownish green. Inhabiting last year’s stems. Type species : P. carduorum Rehm.

18. PYRENOPEZIZA CARDUORUM Rehm in Winter, Diag. u. Notiz, zu Rehm’s Ascom. 562. 1872.

Niptera carduorum (Rehm) Winter in Krypt. FI. 3: 555. 1896. — Ephelina carduorum (Rehm) Rehm in Ber.

bayr. bot. Ges. 13: 183. 1912. — Mollisia carduorum (Rehm) Gremmen in Fungus 25: 5, 1955.

Material: Rehm, Ascom. 68, Pyrenopeziza carduorum Rehm, type (S). Host: Cirsium spec.

19. Pyrenopeziza pastinacae (Nannf.) Gremmen, comb. nov.

Mollisia pastinacae Nannf. in Nova Acta Reg. Soc. Sei. Ups. ser. 4. 8 (2) : 127. 1932 (basionym).

Material: Flora suec. 1906, Mollisia pastinacae Nannf., isotype (UPS). Host: Pastinaca, Anthriscus, Bidens, Cirsium, Heracleum, Angelica.

20. PYRENOPEZIZA URTICICOLA (Phill.) Boud., Disc. d’Eur. 135.1907.

Mollisia urticicola Phill., Man. Brit. Disc. 177. 1887 (basionym).

Material: no. 1109, Mollisia urticicola Phill., teste J. A. Nannfeldt (Grd). Host: Urtica dioica.

21. Pyrenopeziza depressuloides (Gremmen) Gremmen, comb. nov.

Mollisia depressuloides Gremmen in Fungus 25: 9. 1955 (basionym).


Material: no. 807, Mollisia depressuloides Gremmen, type (Grm). Host: Arctium

spec.

22. PYRENOPEZIZA DIGITALINA (Phill.) Sacc., Syll. Disc. 358. 1889.

Mollisia digitalina Phill., Man. Brit. Disc. 190. 1887 (basionym).

Material: no. 1434, Mollisia digitalina Phill. (Grm). Host: Digitalis purpurea.

Sectio 5. Melipeziza Gremmen, sect. nov.

Receptaculum cellulis melleis vel phaeis formatum, margine cellulis elongatis, hyalinis vel subhyalinis. In

caulibus anni praeteriti emortuis. Sectionis species typica: P. leucostoma (Karst.) Nannf.

Receptacle melleous or light brown, its margin with colourless or pale coloured elongate cell-

protuberances. Inhabiting last year’s stems. Type species: P. leucostoma (Karst.) Nannf.

23. PYRENOPEZIZA LEUCOSTOMA (Karst.) Nannf. in Nova Acta Reg. Soc. Sci. Ups. ser. 4. 8 (2) : 153.

1932.

Trochila leucostoma Karst, in Not. Sällsk. Fauna Fl. fenn. 2:245. 1870 (basionym). — Mollisia leucostoma

(Karst.) Karst., Myc. fenn. 1: 203. 1871. — Niptera leucostoma (Karst.) Sacc., Syll. Disc. 483. 1889.

Material: Niptera leucostoma Karst., type (H). Host: Artemisia spec.

24. PYRENOPEZIZA GALII-VERI (Karst.) Sacc., Syll. Disc. 356. 1889.

Mollisia galii-veri Karst., Myc. fenn. 1: 203. 1871 (basionym).

Material: Flora suec. 1988, Pyrenopeziza galii-veri (Karst.), (UPS). Host : Galium verum.

25. PYRENOPEZIZA COMPRESSULA Rehm in Krypt. Fl. 3: 618. 1896. Material: Rehm, Ascom. 69,

Pyrenopeziza compressula Rehm f. 4 spored, type (S). Host : Galeopsis tetrahit.

26. PYRENOPEZIZA CHAMAENERII Nannf. in Svensk bot. Tidskr. 22: 134. 1928. Peziza ebuli Karst., Mon.

Pez. 160. 1869. — Mollisia chamaenerii (Nannf.) Gremmen in Fungus 26: 33. 1956.

Material: Peziza ebuli Karst., type (H). Host : Epilobium spec.

27. PYRENOPEZIZA ERYNGII Fuck., Symb. myc. 294. 1869. Material: Rabenh., Fgi. eur. 1614,

Pyrenopeziza eryngii Fuck. (L). Host : Eryngium campestre.

28. PYRENOPEZIZA SOLIDAGINIS (Karst.) Schrot, in Cohn, Krypt. Fl. Schles. 3 (2): 114. 1893.

Mollisia solidaginis Karst., Rev. Mon. 173. 1885 (basionym). — Peziza subatra Cke. & Peck in 28th Ann. Rep.

N.Y. State Mus. 66-67. 1876· — Mollisia atrata var. megalospora Ell. & Ev., North Am. Fgi. ser. 2, 2625, 1891.

Material: no. 1088, Pyrenopeziza solidaginis (Karst.) Schrot., neotype, (UPS). Host:

Solidago virgaurea.

44 FUNGUS [JAARG.28 No. 1-4

29. PYRENOPEZIZA RUBI (Fr.) Rehm in Krypt. Fl. 3: 611.1896. Excipula rubi Fr., Syst. myc. 2: 190. 1822 (basionym). — Mollisia rubi (Fr.) Karst., Rev. Mon. 136. 1885.

Material: Rabenh., Herb. myc. 2, 305, Peziza phacidioides Fr. (L). Host: Rubus spec.

30. PYRENOPEZIZA MILLEGRANA Boud., Disc. d’Eur. 133. 1907. Mollisia millegrana (Boud.) Nannf. in Nova Acta Reg. Soc. Sei. Ups. ser. 4. 8 (2) : 127. 1932. Material: Flora suec. 4315, Mollisia millegrana (Boud.) Nannf. (UPS). Host: Ulmaria.

31. PYRENOPEZIZA ARCTII (Phill.) Nannf. in Nova Acta Reg. Soc. Sxi. Ups. ser. 4. 8 (2): 142. 1932.

Peziza arctii Phill. in Proc. Bristol Nat. Soc. N.S. 4: 58. 1883 (basionym). — Mollisia arctii (Phill.) Phill.,

Man. Brit. Disc. 183. 1887. — Belonium arctii (Phill.) Sacc., Syll. Disc. 495. 1889. - Belonidium arctii (Phill.)

Massee, Brit. Fung. Fl.4:225· 1895.

Material: Peziza arctii Phill., type (BM). Host: Arctium spec. 32. PYRENOPEZIZA EBULI (Fr.) Sacc., Syll. Disc. 360.1889.

Peziza atrata var. ebuli Fr., Syst. myc. 2: 148. 1822 (basionym). — Mollisia ebuli (Fr.) Karst., Myc. fenn. 1:

202. 1871.

Material: Rehm, Ascom. 656-b, Mollisia ebuli (Fr.) Karst. (S). Host: Sambucus ebulus.

33. PYRENOPEZIZA PLANTAGINIS Fuck., Symb. myc. 294. 1870. Mollisia plantaginis (Fuck.) Phill., Man. Brit. Disc. 183. 1887. — Peziza atrata var. foliicola Desm. in Ann. Sci.

Nat. sér. 2, Bot. 368-369. 1843.

Material: Desm. Crypt. Fr. Sér. 1, 1313, Peziza atrata var. foliicola Desm. (K).

Host : Plantago lanceolata.

34. Pyrenopeziza pulveracea (Fuck.) Gremmen, comb. nov.

Peziza pulveracea Fuck., Symb. myc. 297. 1870 (basionym). — Mollisia pulveracea (Fuck.) Rehm in Krypt. FI.

3: 532. 1896.

Material: Fgi. rhen. 2191, Peziza pulveracea Fuck., type (G). Host: Ulmaria.

35· Pyrenopeziza polygoni (Lasch) Gremmen, comb. nov.

Peziza polygoni Lasch in Klotzsch, Herb. viv. myc., in Rabenh., Herb. myc. 1842 (basionym).

— Mollisia polygoni (Lasch) Rehm in Krypt. Fl. 3: 527. 1896.

Material: Rehm, Ascom. 70, Niptera polygoni Rehm (L). Host: Polygonum amphibium.

36. PYRENOPEZIZA PETIOLARIS (A. & S. ex Fr.) Nannf. in Nova Acta Reg. Soc. Sci. Ups. ser. 4. 8 (2): 158. 1932.

Hysterium petiolare A. & S., Consp. Fung. Nisk. 59. 1805; ex Fr., Syst. myc 2: 593. 1823 (basionym). —

Trochila petiolaris (A. & S. ex Fr.) Rehm in Krypt. Fl. 3: 132. 1896. — Mollisia petiolaris (A. & S. ex Fr.) Gremmen

in Fungus 24: 9. 1954.

Material : no. 9700, Mollisia petiolaris (A. & S. ex Fr.) Gremmen (L). Host: Acer spec.


37. PYRENOPEZIZA MOLLISIOIDES (Sacc. & Br.) Petr, in Ann. myc. 38: 159. 1940.

Phacidium mollisioides Sacc. & Br. in Rev. myc. 7: 210. 1885 (basionym).

Material: Sydow, Myc. germ. 3533, Pyrenopeziza mollisioides (Sacc. & Br.) Petr. (L). Host:

Euphorbia palustris.

38. Pyrenopeziza stellata (Le Gal) Gremmen, comb. nov.

Mollisia stellata Le Gal in Rev. de Myc. 4: 60. 1939 (basionym).

Material: Mollisia stellata Le Gal, type (PC). Host : Dahlia spec.

39. Pyrenopeziza nannfeldtii Petr, in Ann. myc. 38: 368. 1940.

Mollisia nannfeldtii (Petr.) Gremmen in Ber. Schweiz, bot. Ges. 66: 158. 1956.

Material: no. 1221, Mollisia nannfeldtii (Petr.) Gremmen (Grm). Host: Laserpitium, Stier.

Sectio 6. Fimbriaria Gremmen, sect. nov.

Receptaculum cellulis lateralibus minutis, fimbriatis, incoloratis formatum. In caulibus anni praeteriti

emortuis. Sectionis species typica: P. artemisiae (Lasch) Rehm.

Receptacle laterally provided with minute, fimbriate, colourless cell-excrescences. Inhabiting last year’s

stems. Type species: P. artemisiae (Lasch) Rehm.

40. PYRENOPEZIZA ARTEMISIAE (Lasch) Rehm in Krypt. Fl. 3: 616. 1896.

Peziza artemisiae Lasch in Rabenh. Handb. 1: 344. 1844 (basionym). — Mollisia artemisiae (Lasch)

Gremmen in Fungus 25: 2. 1955.

Material: Peziza artemisiae Lasch, type (L). Host: Artemisia spec.

41. Pyrenopeziza clavata (Gremmen) Gremmen, comb. nov.

Mollisia clavata Gremmen in Fungus 24: 5. 1954 (basionym).

Material: Sydow, Myc. germ. 3147, Mollisia minutella (Sacc.) Rehm, type

(L).

Host: Rubus, Epilobium.

42. Pyrenopeziza tormentillae (Vel.) Gremmen, comb. nov.

Tapesia tormentillae Vel., Mon. Disc. 136. 1934 (basionym).

Material: Herb. Velenovský, 149476, Tapesia tormentillae Vel., type (PR). Host:

Tormentilla spec.

43. Pyrenopeziza lycopincola (Rehm) Gremmen, comb. nov.

Mollisia lycopincola Rehm in Hedwigia 27: 166. 1888 (basionym).

Material: Rehm, Ascom. 910, Mollisia lycopincola Rehm, type (S). Host : Lycopus

europaeus.

Sectio 7. Trichantina Gremmen, sect. nov.

Receptaculum in margine et latere pilis septatis, fuscis instructum. In caulibus anni praeteriti emortuis.

Sectionis species typica: P. escharodes (B. & Br.) Rehm.


Receptacle marginally as well as laterally provided with brown, septate hairs. Inhabiting last year’s

stems. Type species : P. escharodes (B. & Br.) Rehm.

This group forms a transition to the genus Tapesia Fuck.

44. PYRENOPEZIZA ESCHARODES (B. & Br.) Rehm in Krypt. Fl. 3: 612. 1896.

Peziza escharodes B. & Br. in Ann. Mag. nat. Hist. ser. 4. 7: 433. 1871 (basionym). — Lachnella escharodes

(B. & Br.) Phill., Man. Brit. Disc. 262. 1887. — Mollisia escharodes (B. & Br.) Gremmen in Fungus 24: 5. 1954.

Material: Peziza escharodes B. & Br., type (BM). Host : Rubus spec. Sectio

8. Hemipirottaea Gremmen, sect. nov.

Apothecia paraphysibus lanceolatis instructa. In caulibus anni praeteriti emortuis. Sectionis species

typica: P. lanceolata (Gremmen) Gremmen.

Apothecia with characteristic lanceolate paraphyses. Inhabiting last year’s stems. Type species: P.

lanceolata (Gremmen) Gremmen.

This section is intermediate between the section Astromatina and the genus Pirottaea Sacc.

45. Pyrenopeziza lanceolata (Gremmen) Gremmen, comb. nov.

Mollisia lanceolata Gremmen in Fungus 26: 35. 1956 (basionym).

Material: no. 690, Mollisia lanceolata Gremmen, type (Grm). Host: Ulmaria.

ACKNOWLEDGEMENT.—Dr R. A. Maas Geesteranus (Leiden) critically revised the manuscript for

which I am very thankful.

REFERENCES

BOUDIER, EM. 1907. Histoire et Classification des Discomycètes d’Europe. Paris. FRIES, E. 1822-1823. Systema mycologicum 2. Lundae. FUCKEL, L. (1869) 1870. Symbolae mycologicae. Wiesbaden. GREMMEN, J. 1952. Het genus Pyrenopeziza. Levenswijze en cultuur van Pyrenopeziza plantaginis Fuck, in Fungus

22: 22-24. ----------- 1954. Taxonomical notes on Mollisiaceous fungi—I. A study on some Dutch species

growing on Rubus stems. In Fungus 24: 1-8. ----------- 1955. Taxonomical notes on Mollisiaceous fungi—II. Some caulicolous Mollisia species

inhabiting various hosts, mainly Compositae. In Fungus 25: 1-12. KARSTEN, P. A. 1871. Mycologia fennica, Pars 1. Discomycetes. In Bidr. Känn. Finl. Nat. Folk

19.

NANNFELDT, J. A. 1932. Studien über die Morphologie und Systematik der nicht-lichenisierten inoperculaten Discomyceten. In Nova Acta Reg. Soc. Sci. Ups. ser. 4. 8 (2).

PHILLIPS, W. 1887. A manual of the British Discomycetes. London. REHM, H. 1896. Die Pilze Deutschlands, Oesterreichs und der Schweiz. In Rabenhorst’s Kryptogamenflora, ed. 2,

1 (3). Leipzig. — ------ 1912. Zur Kenntnis der Discomyceten Deutschlands, Deutsch-Oesterreichs und der Schweiz.

In Ber. bayr. bot. Ges. 13: 102-206.

NOTE COMPLEMENTAIRE A PROPOS DE CYSTODERMA SUPERBUM HUIJSM.

H.S.C. HUIJSMAN


Dans Fungus 26: 39. 1956, j’ai rejeté l’identité de Cystoderma superbum Huijsm. avec C. haematites (Berk. & Br.) Kd & Mblc sensu Kühner & Maire (non sensu orig.) parce que les spores du premier m’étaient apparues non amyloïdes, aussi bien sur sporée fraîche qu’ à l’examen individuel au microscope, alors que les spores de l’autre, précisées dans la littérature comme étant de même taille et forme, étaient signalées amyloïdes.

Depuis j’ai eu l’occasion d’échanger une correspondance avec M. R. Kühner qui, ayant récemment récolté en Haute-Savoie C. haematites sensu Kühner & Maire, a eu l’amabilité d’attirer mon attention sur le fait que la réaction au chloral-iodé était confiné à la plage suprahilaire de la spore.

J’ai procédé à de nouvelles vérifications et ai pu constater qu’effectivement, vue de face, une spore de C. superbum ne montre aucune trace de réaction dans le Melzer tandis que, de profil, le contour de la plage est souvent légèrement accentué. En examinant une tétrade d’extrémité, suivant le conseil de M. Kühner, on voit nettement une croix de substance amyloïde à l’endroit où les quatre spores sont accolées. Cette localisation de la réaction est plus facile à remarquer et

devient nettement plus apparente si on fait en même temps une comparaison avec des tétrades de spores de C. granulosum ou cinnabarinum qui sont absolument insensibles en présence du même réactif.

Cette constatation rapproche singulièrement la plante de Schüpfheim de la plante de Haute-Savoie. Il reste à préciser si une sporée fraîche de mon champignon est réellement insensible au chloral-iodé. Peut-être n’ai je pas porté assez d’attention à ce fait à l’occasion de l’unique examen effectué lors de la récolte, de sorte que l’absence d’une réaction visible à l’oeil nu reste à démontrer.

En attendant, il ne me semble pas inutile de donner une diagnose, légèrement modifiée, de Cystoderma superbum, espèce qui, contrairement à ma première opinion, appartient plutôt à la

section Amianthinae Smith & Singer.

CYSTODERMA SUPERBUM Huijsm.

Chapeau 20-55 mm de large, convexe puis plan-convexe, légèrement granuleux, purpuracé-vineux, à frange

plus pâle; chair assez mince, subconcolore, odeur et saveur pratiquement nulles. Lamelles assez serrées,

arrondies-adnées, crème à saumon; arête érodée. Pied 35-60 x 4-8 mm, d’abord farci d’une moelle blanchâtre,

puis tubuleux, revêtu de squamules granulo-fibrilleuses jusqu’à une zone annulaire filamenteuse, plus pâles

que le fond qui est concolore au chapeau, glabrescent.

Spores blanches en tas, 3,8-4,6 x 2,9-3,3 μ, lisses, ellipsoïdes, non amyloïdes à l’exception de la plage

suprahilaire. Basides à quatre stérigmates. Cystides absentes. Couche superficielle du chapeau à sphérocystes

nombreuses incrustées d’un pigment brun-rougeâtre, plus foncée dans la potasse. Hyphes bouclées.

THE STIPITATE HYDNUMS OF THE NETHERLANDS—III

Phellodon P. Karst, and Bankera Coker & Beers ex Pouz.

R. A. MAAS GEESTERANUS Rijksherbarium, Leiden

A revision of the genera Phellodon and Bankera is given. No new combinations are proposed.

PHELLODON P. Karst.

Phellodon P. Karst, in Rev. mycol. 3 (No. 9): 19. Jan. 1, 1881 & in Medd. Soc. F. Fl. fenn. 6 :15. 1881. — Hydnum [sect.] Phellodon (P. Karst.) J. Schroet. in Cohn, Krypt.-Fl. Schles. 3 (1) : 456. 1888. —Calodon subgen. Phellodon (P. Karst.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 48: 357. 1889. — Type species: Hydnum nigrum Fr. ex Fr., see Donk (1956: 108).

Carpophores terrestrial, stipitate, not fleshy, frequently confluent. Pileus covered with tomentum, anoderm.

Stipe either tomentose, binding plant debris, or glabrescent and clean. Context fibrous, more or less zonate,

almost homogeneous in some species, clearly duplex in others, i.e. soft and spongy without, firm to hard

within. Hymenium covering spines on underside of pileus. Spines some shade of grey at maturity. Basidia

tetrasporous. Spores subglobose, even in outline, minutely spinulose, white in mass. Odour of fenugreek when

dried. Hyphae without clamp connections.

Unlike in Hydnellum and Sarcodon, the hyphae of all species of the present genus lack clamp connections. This condition deprives us of an important means of classing the species in more or less convenient groups. Another inconvenience is in the fact that the KOH-test, used to advantage in the genera mentioned above, gives unstable results with some species of Phellodon. There seems to be no way around a grouping of the species on the basis of purely macroscopical features, which, in view of the similarity of a number of the species, is no easy matter.

Such an arrangement has already been suggested by Coker & Beers, but needs emending in that (i) their Ph. graveolens ( = Ph. melaleucus) seems better placed near Ph. tomentosus, and (ii) Ph. carnosus Banker has to be removed altogether, since this species is a true Bankera (see Snell, Dick, Jackson & Taussig, 1956: 174)·

As far as the European species are concerned, a more precisely worded, although linguistically less accessible, grouping was proposed by Pouzar (1956: 74) who, rather than taking the duplex nature of the context of the entire carpophore as a distinguishing feature, restricted this character to the stipe only. This arrangement is adopted in the key below.

With regard to the American species of the genus, I was fortunate in having had on loan a number of collections from Coker’s herbarium, and been given a part of the type as well as two other collections of Phellodon brunneoroseus Snell, Dick & Jackson. From the study of this material it appears that the following groups may be recognised.

The first group consists of Phellodon aiboniger (Peck) Banker and Ph. niger; the second group of Ph. confluens and Ph. putidus (Atk.) Banker; the third group of Ph. brunneo-olivaceus Coker &

Beers, Ph. brunneoroseus, and Ph. mela-

Nov. 1958] R. A. Maas Geesteranus : Stipitate Hydnums—III 49

leucus; the fourth group of Ph. tomentosus. Of uncertain position are Phellodon cokeri Banker and Ph. ellisianus Banker, the former being tentatively placed in the group of Ph. confluens, the latter in the group of Ph. melaleucus. A fifth group, finally, seems necessary to accomodate Ph. delicatus (Schw.) Banker.

KEY TO THE EUROPEAN SPECIES

I a. Surface of stipe spongy-tomentose ; context of stipe clearly duplex. 2a. Core in stipe black, green in KOH ................................................................................. Ph. tiiger, p. 53 2b. Core in stipe grey-brown, usually not discolouring green in KOH ……………………….. Ph. confluens, p. 49

Ib. Surface of stipe hard, glabrous or nearly so; context of stipe homogeneous. 3a. Pileus except for tomentose margin azonate or colour zones indistinct, without yellowish tints; context of pileus with purplish or slaty hues, green in KOH ………………………………………………… Ph. melaleucus, p. 50 3b. Pileus clearly and densely zonate, with yellowish tints; context of pileus pallid to pale grey-brown, not discolouring green in KOH ............................................................................................ …… Ph. tomentosus, p. 54

PHELLODON CONFLUENS (Pers.) Pouz.

Hydnum * confluens Pers., Mycol. europ. 2: 165. 1825; not Hydnum confluens Peck in Rep. N.Y. State Mus. nat. Hist. 26: 71. 1874 (see remarks). — Phellodon confluens (Pers.) Pouz. in Česka Mykol. 10: 74. 1956. — Type: Hydnum confluens, var. pileis concretis. Prope Clamar” (L 910. 256-1608, here chosen as lectotype).

Hydnum amicum Quél. in Grevillea 8: 115. 1880. — Calodon amicus (Quél.) Quél. in C.R. Assoc, franç. Avanc. Sci. 12: 504. (1883) 1884. — Sarcodon amicus (Quél.) Quél., Ench. Fung. 189. 1886. — Phellodon amicus (Quél.) Banker in Mycologia 5: 62. 1913. — Hydnellum amicum (Quél.) Ragab in Mycologia 45: 944. 1953. — Type: (in absence of material, information Prof. A. Maublanc) represented by Quél. in Grevillea 8: pl. 131 fig. 1. 1880.

Hydnum vellereum Peck in Ann. Rep. N.Y. State Mus. 50 : no. 1897. — Phellodon vellereus (Peck) Banker in Mem. Torrey bot. Cl. 12: 168. 1906. — Type: not seen (NYS).

MISAPPLICATION: Hydnum cinereum Bull, ex Fr. sensu Fr., Epier. Syst. mycol. 508. 1838; Hym. europ. 604. 1874; not Hydnum cinereum Bull, ex Fr., Syst. mycol. 1: 404. 1821 — Bankera violascens, which see for discussion; not Hydnum cinereum (Batsch) ex Pers., Mycol. europ. 2: 168. 1825 = Phellodon niger.

DESCRIPTIONS.—Banker in Mem. Torrey bot. Cl. 12: 168. 1906 (Phellodon vellereus); Boudier, Icon, mycol. 4: 86. 1911 (Hydnum amicum); Bourdot & Galzin, Hym. France 462. 1928 (Calodon amicus); Coker & Beers, Stip. Hydn. east. U.S. 23. 1951 (Phellodon amicus); Donk in Med. Nederl. mycol. Ver. 22: 48. 1933 (Phellodon amicus).

ILLUSTRATIONS.—Boudier, Icon, mycol. 1: pl. 167. 1904-1910 (Hydnum amicum; dark brown centre rather unusual, otherwise good) ; Coker & Beers, Stip. Hydn. east. U.S. pl. 15. 1951 (Phellodon amicus; photogr.) ; Fl. batava 20: pl. 1540. 1898 (Hydnum cinereum; unrecognisable) ; Patouillard, Tab. anal. Fung. 3 : fig. 246. 1884 (Hydnum amicum; passable) ; Quélet in Grevillea 8 : pl. 131 fig. I. 1880 (Hydnum amicum; mediocre) & in C.R. Assoc, franç. Avanc. Sci. 12 : pl. 6 fig. 14. (1883) 1884 (Calodon amicus; same as preceding, but colours different) ; Rolland, Atl. Champ. France pl. 101 fig. 223. 1910 {Hydnum amicum ; colours too bright and contrasting).

DIAGNOSTIC CHARACTERS.—Carpophores solitary or concrescent, often several fusing into complex groups.

Pileus plane to depressed, surface thickly woolly-tomentose, more or less uneven, becoming matted or pitted

from collapsing tomentum, sometimes in places also radially ridged, without concentrical zones, whitish when

young, from centre outwards turning various shades of grey-brown (rarely very dark), at times suffused with

faint lilaceous tint, margin long remaining whitish tomentose. Stipe stocky, equal or


tapering downwards, sometimes extremely short, thick-tomentose, becoming matted and somewhat shiny,

concolorous with cap or darker, often obscured by adhering vegetable debris. Spines decurrent, finally silvery

grey, sometimes distinctly suffused with lilac. Context thick, duplex, very firm in central part of stipe, zonate,

grey-brown to bistre, not discolouring in KOH or giving a transient dingy olive green colour. Odour of

fenugreek when dry.

HABITAT.—On acid, humous soil in frondose woods, the record of the vicinity of Pinus probably being incidental.

DISTRIBUTION.—In the central, southern, and eastern parts of the country, not uncommon. ILLUSTRATIVE COLLECTIONS.—Utrecht: Zeist, Wulperhorst, 30 VIII 1954, C. Bas 562 (L);

Zeeland: Zeeuws Vlaanderen, Hulst, 12 X 1951, B.J.J. R. Walrecht (L). AUTHENTIC COLLECTION.—Calodon amicum Q., Herb. Bourdot 14710, Champignons de l’Allier,

bois entre les Robinets et la Ronde; Iseure. Leg. H. B. 2230, 6 IX 1897. “Calodon amicum Quélet!” in vivo determinavit Quélet n. 150! (PC).

EXSICCATI.—Brinkmann, Westfäl. Pilze, Lief. 2: 95 (Hydnum candicans; K) ; Fuckel, Fungi rhen. 1344 (Hydnum tomentosum; K) ; Kryptog. exs. vindob. 3014 (Hydnum amicum; C, K, L, PR) ; Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. 1401 (Hydnum amicum; PR, UPS); 1402 (Hydnum amicum; PR); Roumeguère, Fungi gall. exs. 2313 (Hydnum nigrum; K) ; Saccardo, Mycoth. ven. 825 (Hydnum cinereum; K) ; von Thiimen, Mycoth. univers. 1907

(Hydnum amicum; C, L).

According to the original description Hydnum confluens of Peck is different from Persoon’s species. Unfortunately, the type material failed to reach me in time, but the description of the pileus as “thin... zonate, slightly fibrous-tomentose, hygrophanous, dark brown when moist...”

makes it improbable that the species should be identified with Phellodon confluens.

PHELLODON MELALEUCUS (Fr. ex Fr.) P. Karst.

Hydnum melaleucum Fr., Obs. mycol. I: 141. 1815; ex Fr., Syst. mycol. 1: 406. 1821; Elench. Fung. 1: 131. 1828; Epicr. Syst. mycol. 510. 1838; Hym. europ. 606. 1874. —Hydnellum melaleucum (Fr. exFr.) P. Karst, in Medd. Soc. F. Fl. fenn. 5: 41. 1879. —Phellodon melaleucus (Fr. ex Fr.) P. Karst, in Rev. mycol. 3 (No. 9) : 19. Jan. 1, 1881 & in Medd. Soc. F. Fl. fenn. 6: 15. 1881 —Calodon melaleucus (Fr. ex Fr.) Quél., Ench. Fung. 191. 1886. — Calodon graveolens var. melaleucus (Fr. ex Fr.) Quél., Fl. mycol. 445. 1888. — Hydnum graveolens subsp. melaleucum (Fr. ex Fr.) Légué, Catal. rais. Basidiomyc. Mondoubleau 158. 1908. — Type: non-existing, cf. Banker in Mycologia 5: 63. 1913. — Type locality: Sweden (“In pinetis, Smolandiae, V. Gothiae in Râda âs, prope Holmiam.”—Lindblad, Syn. Fung. Hydn. Suec. nasc. 12. 1853).

Hydnum pullum Schaeff., Fung. Icon. 4: 98. 1774; not Hydnum pullum Sw. in Svensk Vet Akad. nya Handl. 31: 248. 1810 = Phellodon niger. — Hydnum leptopus var. ß pullum (Schaeff.) ex Pers., Mycol. europ. 2: 171. 1825. — Hydnum pullum (Schaeff. ex Pers.) Duby, Bot. Gall. 2: 776. 1830. — Calodon pullus (Schaeff. ex Pers.) Quél. in C.R. Assoc, franç. Avanc. Sci. 22: 488, (5 of reprint). (1893) 1894. — Phellodon pullus (Schaeff. ex Pers.) Banker in Mycologia 5: 62. 1913. — Type: represented by Schaeff., Fung. Icon. 3: pl. 272. 1770.

Hydnum tomentosum var. atro-album Alb. & Schw., Consp. Fung. 266. 1805; ex Pers., Mycol. europ. 2: 167. 1825. — Type: not known to be in existence. — Type locality: Germany, Oberlausitz “Moholzer Haide; Seer Busch” (Alb. & Schw., l.c. 267).

Hydnum leptopus var. γ graveolens Pers., Mycol. europ. 2: 171. 1825. — Hydnum graveolens. (Vers.) Fr., Epicr. Syst. mycol. 509. 1838 (“Delast. ! in litt, primus inventor!”) ; Hym. europ. 605. 1874.

Nov. 1958] R. A. Maas Geesteranus: Stipitate Hydnums—III 51

— Hydnellum graveolens (Pers.) P. Karst, in Medd. Soc. F. Fl. fenn. 5: 41. 1879. — Phellodon graveolens (Pers.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 37: 96. 1882. — Calodon graveolens (Pers.) Quel., Ench. Fung. 191. 1886. —Phellodon melaleucus f. graveolens (Pers.) Nikolajeva in Pl. cryptog. 9: 488. 1954 (“Donk”).—Type: “Hydnum graveolens. Mycol. Europ. S. 3. p. 171. Gallia” [misit Delastre] (L 910.263-874, here chosen as lectotype).

Hydnum fuscum foetens Seer., Mycogr. suisse 2 : 519. 1833. — T ype : not known to be in existence. — Type locality: Switzerland, Sauvabelin.

Calodon graveolens f. nigricans Bourd. & Galz. in Bull. Soc. mycol. France 40: 119. 1924; Flym. France 462. 1928. — Type: not known to be in existence. ·— Type locality: France.

Calodon graveolens f. ramosus Bourd. & Galz. in Bull. Soc. mycol. France 40: 119. 1924; Hym. France 462. 1928. — Type: not known to be in existence. -— Type locality: France.

Phellodon melaleucus f. major Donk in Med. Nederl. mycol. Ver. 22: 50. 1933. — Type: not indicated, presumably Hydnum melaleucum, Gelderland: Lochem, Boekhorst, 30 VIII 1878, J. Staring (GRO).

Phellodon melaleucus f. violascens Donk in Med. Nederl. mycol. Ver. 22: 50. 1933. — Type: Phellodon melaleucus f. violascens, Netherlands, locality unknown, probably Meyendel near Wassenaar, X 1930, (Herb. Donk 2458).

DESCRIPTIONS.—Banker in Mem. Torrey bot. Cl. 12: 169. 1906 (Phellodon graveolens); Bourdot & Galzin, Hym. France 462. 1928 (Calodon graveolens); Coker & Beers, Stip. Hydn. east. U.S. 26.

1951 (Phellodon graveolens) ; Donk in Med. Nederl. mycol. Ver. 22: 49. 1933; Konrad & Maublanc, Icon. sel. Fung. 5 : text to pl. 477 fig. 1. 1932 (Calodon graveolens).

ILLUSTRATIONS.—Coker in J. Mitchell sci. Soc. 41: pl. 57. 1926 (Phellodon graveolens; photogr.); Coker & Beers, Stip. Hydn. east. U.S. pl. 18 (below). 1951 (Phellodon graveolens; photogr.); FI. batava 19: pl. 1480. 1893 (Hydnum; good) ; Fries, Icon. sel. Hym. 1 : pl. 6 fig. 1. 1867 (Hydnum graveolens; passable) ; Konrad & Maublanc, Icon. sel. Fung. 5: pl. 477 fig. 1. 1932 (Calodon graveolens ; passable); Michael-Schulz., Führer Pilzfr. 3: pl. 309. 1927 (Hydnum; good); Nikolajeva in Pl. cryptog. 9: fig. 11. 1954 (Phellodon melaleucus f. melaleucus ; passable); Pouzar in Ceska Mykol. 10: fig. 6 & 7. 1956 (photogr.); Schaeffer, Fung. Icon. 3: pl. 272. 1770 (Hydnum pullum; good excepting colour of context of stipe in fig. 4).

DIAGNOSTIC CHARACTERS.—Carpophores usually confluent, sometimes fused into complex mass of partly

imbricate pilei. Pileus plane with depressed to umbilicate centre, or infundibuliform, orbicular to flabelliform,

surface thinly felted-tomentose, very soon glabrescent, becoming comparitively even with only centre pitted

and radially fibrillose-striate to ridged further outwards, or harshly pitted, lamellate-scaly, and jaggedly as-

perate all over, frequently with a few concentrical corrugations, shining, with obscure concentrical colour zones

or azonate, grey-brown to black-brown with or without purplish hue, or blackish olive, darkest in centre, with

the margin long remaining white- or creamy-tomentose, more rarely lilaceous. Stipe central to lateral, very

slender to stocky, tapering downwards to equal, rooting or springing from brown-felted subiculum, fibrillose to

glabrous, usually free from adhering vegetable debris, concolorous with pileus to nearly black. Spines

decurrent, finally ashy grey, at times suffused with lilaceous hue. Context thin, hardly duplex in cap,

homogeneous in stipe, soft throughout or with somewhat firmer zone just below surface at junction of pileus

and stipe, indistinctly zonate, grey-brown with purplish or slaty tinge in pileus, turning olive green in KOH,

fuscescent with age, bistre and not discolouring in KOH in stipe. Odour of fenugreek when dry, but of varying

intensity.

HABITAT.—On acid, sandy soil in both frondose and coniferous woods. DISTRIBUTION.—In most parts of the country, fairly common.


ILLUSTRATIVE COLLECTIONS.—Drente: Dwingelo, Lheederzand, 12 X 1953, C. Bas (L); Utrecht: Baarn, Soestdijk, 15 IX 1957, G. A. de Vries (L).

EXSICCATI.—Brinkmann, Westfäl. Pilze, Lief. 2: 97 (Hydnum; K, L) ; Fuckel, Fungi rhen. 1346 (Hydnum; K) ; Karsten, Fungi fenn. exs. 246, 907 (Hydnum tomentosum; K) ; Kavina & Hilitzer,

Cryptog. cechoslov. exs. 251 (Hydnum; PR) ; Klotzsch, Herb, vivum mycol. 122 (Hydnum; L, PR) ; Klotzsch, Herb, mycol., ed. nova: 415 (Hydnum; K) ; Kryptog. exs. vindob. 2109 (Hydnum gra-veolens; C, K, L, PR, the latter uncertain); Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. 351 (Hydnum; C, PR); 1403 (Hydnum; PR); Oudemans, Fungi neerl. exs. 234 [Hydnum; K) ; Petrak, Fl. Bohem. Morav. exs. ser. 2 (1) : 1361 (Hydnum graveolens; PR); Rabenhorst, Fungi europ. 1104b (Hydnum graveolens; L) ; Rabenhorst, Herb, mycol. ed. 2: 415 (Hydnum; L, PR) ; Roumeguère, Fungi gall. exs. 2312 (Hydnum; BR); Sydow, Mycoth. germ. 1052 (Hydnum graveolens; K, L, PC) ; Sydow, Mycoth. march. 908 (Hydnum graveolens; K) ; von Thümen, Fungi austr. 827 (Hydnum; PR).

While it is true that the extreme variability of the present species has been an impediment to its delimitation, I believe that the value attached by later authors to Fries’s description of Hydnum graveolens (1838: 509) only contributed to the confusion. Apparently, it escaped notice that it is not Fries’s description that matters, but Persoon’s. Fries, in citing “H. leptopus γ. Pers. M. E. 2 p. 171,” only made a recombination, raising the original variety to a specific level. So, for a clear understanding of what was meant by ‘graveolens’, the description by Persoon and the material in his herbarium ought to be consulted.

In Herb. Persoon there are three sheets containing material that was sent in by Delastre, all representing Phellodon melaleucus. One is labelled ‘hydnum fragrans’, and is obviously written in the collector’s jaunty hand. Incidentally, Lloyd, who has seen the material, made a note on the cover of the sheet, indicating the specimens as “the type of leptopus var. graveolens Pers or Hydnum graveolens Fr ,” as he found them better developed. However, as the sheet lacks any comment in Persoon’s hand which would provide evidence of the material having been used for

the description, the specimens had better be left out of consideration. Of the two other sheets one has no longer the original label. Since, moreover, the specimens

are glued to a piece of paper differing in texture from that of the other sheet, there is no knowing whether the specimens of the two sheets really belonged to one and the same collection. It seems advisable to leave the specimens of this disputable sheet out of consideration as well.

The last sheet, finally, has attached to it a label written by Persoon, mentioning the wrong part of ‘Mycol. europ.’ (S. 3), but the correct page (p. 171). I consider this sheet to contain the lectotype, but regrettably enough the specimens are inferior to those of the other sheets.

For the description of this (and the other) material I may refer to Donk (1933: 50) under forma 2. However, as already observed by this author, he did not believe "dass es sich dabei um mehr oder weniger natürliche Ausschnitte handelt.” One may, as a matter of fact, find every conceivable form intermediate between ‘f. melaleucus’ and ‘f. graveolens’. Since, moreover, specimens


of the same collection may be seen to vary in either direction, an infra-specific segregation seems superfluous.

In this country the majority of the specimens may be referred to what Donk described under forma 2 (Hydnum graveolens). These are equally often found in coniferous as in frondose woods.

As to the more heavily built type, forma 1 (Hydnum melaleucum) of Donk, there seems to be a slight preference for frondose woods, but once again it is emphasised that, owing to the variability of the species, it is hard to be definite in one’s statements.

Blytt (1905: 131) described a new species, Hydnum myriopedum, of which, unfortunately, no material could be traced in the Botanical Museum, Oslo. From the description (translated: “... internally the pileus is indistinctly zonate... the spines are grey-white. The fungus is leathery...”) it can be gathered that a Phellodon is meant, although the species is placed among the Hydnellums. The words describing the pileus as “but otherwise without zones”, as well as the habitat, “in birch-wood”, exclude Phellodon tomentosus. While the colours of the pileus, “rust-coloured, later on black-brown...” are not good of Phellodon confluens, the description of the carpophores as “confluent to a large, many-but short-stemmed, seemingly sessile cake...” seems equally unsuitable to designate Ph. niger. The only European species left is Ph. melaleucus, and I would not doubt to identify H. myriopedum with this species if it were not for the rusty colour of

the cap.

PHELLODON NIGER (Fr. ex Fr.) P. Karst.

Hydnum nigrum Fr., Obs. mycol. I: 134. 181 5 ; ex Fr., Syst. mycol. 1:404. 1821 ; Elench. Fung. 1: 131. 1828 ; Epicr. Syst. mycol. 509. 1838 ; Hym. europ. 605. 1874. — Hydnellum nigrum (Fr. ex Fr.) P. Karst, in Medd. Soc. F. Fl. fenn. 5: 41. 1879. —Phellodon niger (Fr. ex Fr.) P. Karst, in Rev. mycol. 3 (No. 9) : 19. Jan. 1, 1881 & in Medd. Soc. F. Fl. fenn. 6: 15. 1881. — Calodon niger (Fr. ex Fr.) Quél., Ench. Fung. 191. 1886. — Type: non-existing, cf. Banker in Mycologia 5: 62. 1913. — Type locality: Sweden (“In pinetis passim, Smolandiae, Ostrogothiae, Uplandiae etc.”—Lindblad, Syn. Fung. Hydn. Suec. nasc. 11. 1853).

Hydnum suberosum varietas γ cinerea Batsch, Elench. Fung. Cont. 2: 107. 1789. — Hydnum cinereum (Batsch) ex Pers., Mycol. europ. 2: 168. 1825; not Hydnum cinereum Bull, ex Fr., Syst. mycol. 1:404. 1821 = Bankera violascens, which see for discussion ; nor Hydnum cinereum Bull, ex Fr. sensu Fr., Epicr. Syst. mycol. 508. 1838; Hym. europ. 604. 1874 = Phellodon confluens.— Hydnum pullum Sw. in Svensk VetAkad. nya Handl. 31: 248. 1810 (name change); not Hydnum pullum (Schaeff. ex Pers.) Duby = Phellodon melaleucus. — Hydnum olivaceo-nigrum Seer., Mycogr. suisse 2: 520. 1833 (name change). — Hydnum nigrum var. olivaceo-cinereum Sacc. in Michelia 1: 4. 1877 (non vidi) ; in Fl. ital. cryptog. 1 (15) : 1093. 1916 (“Fr.”) (Hydnum nigrum b. pileo olivaceo-cinereo, Fr., Syst. mycol. 1: 404. 1821]. — Hydnum melilolinum Quél. in Bull. Soc. bot. France 25: 290. 1879 (name change). —Calodon niger var. melilotinus (Quél.) Quél., Ench. Fung. 191. 1886; FI. mycol. 444. 1888. — Hydnum nigrum subsp. *H. melilotinum (Quél.) Sacc., Syll. Fung. 6: 442. 1888. — Phellodon niger f. melilotinus (Quél.) Donk in Med. Nederl. mycol. Ver. 22 : 47. 1933· — Hydnum nigrum var. melilotinum (Quél.) Lundell in Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. Fasc. 29-30: 3. 1947. — Type: represented by Batsch, Elench. Fung. Cont. 2: pl. 40 fig. 223. 1789.

DESCRIPTIONS.—Banker in Mem. Torrey bot. Cl. 12: 166. 1906; Bourdot & Galzin, Hym. France

461. 1928 (Calodon niger & var. melilotinus); Donk in Med. Nederl. mycol. Ver. 22: 46. 1933; Konrad & Maublanc, Icon. sel. Fung. 5: text to pl. 476. 1927 (Calodon niger).

ILLUSTRATIONS.—Batsch, Elench. Fung. Cont. 2 : pl. 40 fig. 223. 1789 (Hyd-


num suberosum var. γ cinerea; good); Fl. batava 25: pi. 1980b. 1920 (Hydnum; passable); Fries, Icon. sel. Hym. 1: pl. 5 fig. 2. 1867 (Hydnum; mediocre); Gillet, Champ. France pl. 321.1878-1890 (Hydnum; fairly good except context) ; Konrad & Maublanc, Icon. sel. Fung. 5: pl. 476. 1927 (Calodon; fairly good except context); Nikolajeva in PL cryptog. 9: fig. 10. 1954 (passable); Pa-

touillard, Tab. anal. Fung. 7: fig. 678. 1889 (Calodon; passable); Pouzar in Českâ Mykol. 10: fig. 5. 1956 (photogr.).

DIAGNOSTIC CHARACTERS.—Carpophores often confluent. Pileus obconical to planoconvex, becoming depressed

or infundibuliform, surface thickly tomentose, even to colliculose, glabrescent from collapsing tomentum,

becoming pitted or lamellate in centre and radiately striate or ridged further outwards, or pitted all over,

occasionally with concentrical corrugations, somewhat shining, with indistinct concentrical colour zones or

azonate, pale bluish grey when very young, at times distinctly violaceous, soon from centre outwards turning

uniformly bluish black to greyish olive, greyish brown when dried, margin whitish. Stipe slender to stocky,

ventricose or with bulbous base from thick spongiose covering, surface becoming matted, fuscous to black, with

tomentum some tint of olive-grey, usually covered by and enclosing mosses and plant detritus. Spines

decurrent, finally ashy grey. Context thick, thinning with age, distinctly duplex both in cap and stipe, outer

parts soft, spongy, concolorous with surface, core in stipe and harder parts above spines in pileus corky to

woody, very little zonate in pileus, bluish black to black, turning olive green to blue-green in KOH. Odour of

fenugreek when dry.

Habitat.—On leaf-humus or acid sandy soil in frondose or, more rarely, mixed woods. Distribution.—Known from southern to eastern parts of the country, but nowhere common. Il lustrative col lections.—Overijsel: Denekamp, Borgbos, 1 X 1950, Maas G. 7506 (L) ;

Utrecht: Baarn, Pijnenburg, 3 X 1954, G. A. de Vries (L). Exsiccati.—Allescher & Schnabl, Fungi bavar. 132 (Hydnum; L) ; Fuckel, Enum. Fung.

Nassov., ser. 1: 947 (Hydnum; L); Fuckel, Fungi rhen. 1347 (Hydnum; K) ; Karsten, Fungi fenn. exs. 908 (Hydnum; K) ; Kryptog. exs. vindob. 2016 (Hydnum; C, K, L, PC, PR); Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. 350 (Hydnum; C, PR) ; 1404 (Hydnum nigrum var. melilotinum; PR) ; Rabenhorst, Fungi europ. 1004 (Hydnum graveolens; K, L) ; 1307 (Hydnum; K, L) ; Roumeguère, Fungi gall. exs. 2007 (Hydnum graveolens; L) ; 2313 (Hydnum; BR, L) ;

Roumeguère, Fungi sel. exs. (K).

Phellodon tomentosus (L. ex Fr.) Banker

Hydnum tomentosum L., Sp. PI. 2: 1178. 1753; Fl. suec., ed. 2, 455. 1755; ex Fr. Syst. mycol. 1 : 405. 1821; Elench. Fung. 1: 131. 1828 (erroneously as H. subtomentosum) ; Epicr. Syst. mycol. 510. 1838. —Phellodon tomentosus (L. ex Fr.) Banker in Mem. Torrey bot. Cl. 12: 171. 1906. — Calodon tomentosus (L. ex Fr.) R. Maire in Publ. Inst. bot. Barcelona 3 (4): 36. 1937. — Type: non-existing, cf. Banker in Mycologia 5: 64. 1913. — Type locality: Sweden, “in sylvis acerosis Dalekarliae” (Linn., Fl. suec. 383, no. 1099. 1745).

Hydnum cyathiforme Schaeff., Fung. Icon. 4: 93. 1774; ex St.-Amans, Fl. agen. 545. Apr. 1821 ; not Hydnum cyathiforme Fr., Syst. mycol. 1: 405. 1821 = Hydnellum velutinum var. scrobiculatum & var. zonatum. — Hydnellum cyathiforme (Schaeff. ex St.-Amans) P. Karst, in Medd. Soc. F. Fl. fenn. 5: 41. 1879. —Phellodon cyathiformis (Schaeff. ex St.-Amans) P. Karst, in Rev. mycol. 3 (No. 9) : 19. Jan. 1, 1881 & in Medd. Soc. F. Fl. fenn. 6: 15. 1881. — Calodon cyathiformis (Schaeff. ex St.-Amans) Quél., Ench. Fung. 191. 1886 — Type: (non-existing, cf. Banker in Mycologia 5: 64. 1913) represented by Schaeffer, Fung. Icon. 2: pl. 139. 1763.


Hydnum zonatum Batsch, Elench. Fung. m. 1783; not Hydnum zonatum sensu Batsch, Elench. Fung. Gont. 2: 109. 1789 = Hydnellum velutinum var. zonatum, cf. Maas Geesteranus in Fungus 27 : 69. 1957. -—- Type: represented by Schaeffer, Fung. Icon. 2: pl. 139. 1763 (selected).

Hydnum leptopus Pers., Mycol. europ. 2: 170. 1825 (excluding var. ß & var. γ = Phellodon melaleucus). — Type: “Hydnum leptopus var. a. Mycol. Eur. 1. p. 170. Bois de Sapin au haut du Jura” (L. 910. 263-1317).

Hydnum coriaceo-membranaceum Schw. in Trans. Amer. phil. Soc. 4: 162. 1834.—Phellodon coriaceo-membranaceus (Schw.) Banker in Mem. Torrey bot. Cl. 12: 172. 1906. — Type: destroyed, cf. Banker, l.c. 173. — Type locality: U.S.A., Pennsylvania, Bethlehem.

Hydnum variecolor Seer., Mycogr. suisse 2: 523. 1833. — Calodon variecolor (Seer.) Quél. in C.R. Assoc. franç. Avanc. Sci. 11: 400 ( 14 of reprint). 1882 (“variicolor”). —Calodon cyathiformis var. variecolor (Seer.) Quél., Fl. mycol. 445. 1888. — Type: not known to be in existence. — Type locality: Switzerland, “Au bois d’Écublens” (Secretan, l.c.).

Hydnum candicans Fr., Epicr. Syst. mycol. 510. 1838; Hym. europ. 606. 1874. — Calodon cyathiformis var. candicans (Fr.) Quél., Ench. Fung. 191. 1886; Fl. mycol. 445. 1888.—Hydnum graveolens var. candicans (Fr.) Bres., Icon, mycol. 22: text to pl. 1057. 1932. — Phellodon melaleucus f. candicans (Fr.) Nikolajeva in Pl. cryptog. 9: 489. 1954 (“Donk”). — Type: represented by Krombholz, Naturgetr. Abb. Beschr. essb. Schw. 1: pl. 5 fig. 12. 1831 (“Hydnum tomentosum”).

Hydnum cyathiforme Schaeff. var. obscurum Sacc. in Michelia 1: 363. 1878 (non vidi); Syll Fung. 6: 443. 1888. — Type: not known to be in existence. — Type locality: “In muscosis udis-silvae Montello Treviso Ital. bor.” (Sacc., l.c. 444).

Hydnum graveolens var. sub zonatum Peck in Bull. N.Y. St. Mus. 75: 24. 1904. — Hydnum subzonatum (Peck) Lloyd, Mycol. Writ. 7: 1339. 1925. — Type: not seen (NYS).

Descriptions.—Banker in Mem. Torrey bot. Cl. 12:171,172.1906 (Phellodon tomentosus & Ph. coriaceo-membranaceus) ; Bourdot & Galzin, Hym. France 462. 1928 (Calodon cyathiformis)·, Coker & Beers, Stip. Hydn. east. U.S. 31. 1951; Donk in Med. Nederl. mycol. Ver. 22: 47. 1933; Konrad & Maublanc, Icon. sel. Fung. 5 : text to pl. 477 fig. 2. 1932 (Calodon cyathiformis).

Il lustrations.—Bresadola, Icon, mycol. 22: pl. 1057 (Hydnum graveolens var. candicans) & 1058 (Hydnum cyathiforme). 1932 (passable) ; Coker & Beers, Stip. Hydn. east. U.S. pl. 20. 1951 (photogr.); Harzer, Naturgetr. Abb. essb. gift, verdächt. Pilze pl. 3a. 1842 (Hydnum; mediocre) ; Konrad & Maublanc, Icon. sel. Fung. 5 : pl. 477 fig. 2. 1932 (Calodon cyathiformis; good) ; Krombholz, Naturgetr. Abb. Beschr. essb. Schw. 1: pl. 5 fig. 12. 1831 (Hydnum; mediocre, stipe misleading); Lloyd, Mycol. Writ. 7: pl. 325 fig. 3115. 1925 [Hydnum subzonatum; photogr.); Michael-Schulz, Führer Pilzfr. 3: pl. 308. 1927 (Hydnum cyathiforme; very good); Nikolajeva in PI. cryptog. 9: fig. 12 (Phellodon melaleucus f. candicans), fig. 13. 1954 (passable); Pouzar in Česka Mykol. 10: fig. 4. 1956 (photogr.); Schaeffer, Fung. Icon. 2: pl. 139. 1763 (Hydnum cyathiforme; excellent).

DIAGNOSTIC CHARACTERS.—Carpophores confluent. Pileus plane with depressed to umbilicate centre, surface

felted-tomentose, very soon glabrescent, becoming pitted-strigose to lamellate-scaly in centre, concentrically

corrugated and radiately fibrillose-striate to ridged further outwards, somewhat shining, isabelline, paler or

darker yellow-brown, or rufous yellow-brown, at times suffused with lilaceous hue, concentrically zoned with

alternating paler and darker bands, margin long remaining white-tomentose. Stipe often but not always arising

from more or less extensive felty yellowish brown subiculum which, while binding vegetable matter, resembles

a bulbous base, slender, equal or tapering downwards, subtomentose but soon turning fibrillose to glabrous

from collapsing tomentum, somewhat shining, concolorous with pileus, darker near base. Spines decurrent,

finally pale brownish grey. Context thin, hardly duplex in pileus, homogeneous in stipe, soft in pileus,

somewhat firmer above spines and in stipe, zonate in latter,.


pallid to grey-brown, darkest in stipe, not discolouring in KOH. Odour of fenugreek when dry, but of varying

intensity.

Habitat.—On acid, sandy soil, without exception in coniferous woods.

Distribution.—From southern to eastern parts of the country, not uncommon. Illustrative col lection.—Gelderland: Hierden, Hulshorster Zand, 14 X 1956, Maas G. 11832

(L). Exsiccati,—Brinkmann, Westfäl. Pilze, Lief. 2: 98 (Hydnum cyathiforme; K, L) ; Ellis, North

Amer. Fungi 927 (Hydnum; L) ; Fuckel, Enum. Fung. Nassov., ser. 1: 945 (Hydnum; L); Fuckel, Fungi rhen. 1345 (Hydnum graveolens?; K) ; Klotzsch, Herb. viv. mycol. 123 (Hydnum; L, PR) ; Kryptog. exs. vindob. 2614 (Hydnum cyathiforme ; C, K, L) ; Litschauer & Lohwag, Fungi sel. exs. europ. 13 (Calodon cyathiformis; Herb. Donk, PR); Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. 69 (Hydnum; C, PR) ; Oudemans, Fungi neerl. exs. 113 (Hydnum cyathiforme; K); Rabenhorst, Fungi europ. 611, 611b (Hydnum; L) ; 2304 (Hydnum; K, L) ; Roumeguère, Fungi gall. exs. 2306 (Hydnum cyathiforme; BR, K); Sydow, Mycoth. germ. 655 (Hydnum cyathiforme; C, K, L, PC); 2304 (Hydnum cyathiforme; C, L, PR); Sydow, Mycoth. march. 206, ion (Hydnum cyathiforme; K); 812 (Hydnum melaleucum; K); von Thümen, Fungi austr. 321 (Hydnum; PR) ; von Thümen, Mycoth. univ. 207 (Hydnum; K, PR) ; 1907 (Hydnum amicum; K,

PR; uncertain).

Bankera Coker & Beers ex Pouz.

Bankera Coker & Beers, Stip. Hydn. east. U.S. 33. 1951 (no Latin description); ex Pouz. in Českâ Mykol. 9: 95. 1955. — Type species: Hydnum fuligineo-album Schmidt ex Fr.

Carpophores terresfrial, stipitate, fleshy, not or little concrescent. Pileus covered with tomentum which on

collapse turns into pellicle. Stipe thinly felted, glabrescent. Context not zonate, homogeneous. Hymenium

covering spines on underside of pileus. Spines some shade of grey at maturity. Basidia tetrasporous. Spores

subglobose, finely tuberculate, white in mass. Odour of fenugreek when dried. Hyphae without clamp

connections.

The present genus has some important characteristics in common with Phellodon: (i) the

spores are white in the mass and not roughly tuberculate; (ii) the hyphae lack clamp connections; (iii) the odour of the dried specimens, although variable in strength, is of fenugreek. These would seem to point to a close affinity of Bankera and Phellodon, constituting a group quite distinct from the genera Sarcodon and Hydnellum which belong to the Thelephoraceae.

KEY TO THE EUROPEAN SPECIES

I a. Carpophores usually solitary. Pileus pale flesh-coloured yellow-brown when young, with tomentum turning into glabrous pellicle, often covered with dirt and vegetable debris B. fuligineo-alba, p. 56

Ib. Carpophores often somewhat concrescent. Pileus pale lilaceous grey when young, with pellicle rupturing into scales, clean .................................................................................................................... B. violascens, p. 58

Bankera fuligineo-alba (Schmidt ex Fr.) Pouz.

Hydnum fuligineo-album Schmidt in Kunze & Schmidt, Mykol. Hefte X: 88. 1817; ex Fr., Syst. mycol. 1: 400. 1821; Epicr. Syst. mycol. 506. 1838; Icon. sel. Hym. 1: 5. 1867; Hym. europ. 601. 1874. — Tyrodon fuligineo-albus (Schmidt ex Fr.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 37:

Nov. 1958] R. A. Maas Geesteranus : Stipitate Hydnums—III 57

91. 1882. — Sarcodon fuligineo-albus (Schmidt ex Fr.) Quél., Ench. Fung. 189. 1886. — Sarcodon violascens var. fuligineo-albus (Schmidt ex Fr.) Quél., Fl. mycol. 447. 1888. - Bankera fuligineo-alba (Schmidt ex Fr.) Coker & Beers, Sdp. Hydn. east. U.S. 34. 1951 (not validly published).— Bankera fuligineo-alba (Schmidt ex Fr.) Pouz. in Česka Mykol. 9: 96. 1955. — Type: probably not in existence. — Type locality: Germany, “... in Fichtenwaldungen bey Bernstadt (im Nonnenwalde) und bey Kauffungen in Sachsen.” (Schmidt, l.c.).

Hydnum fragile Fr. in Ofvers. svensk VetAkad. Förh. 8: 53. 1851 ; Sverig. ätl. gift. Svamp. 51. 1860; Monogr. Hym. 2: 275. 1863; Hym. europ. 599. 1874; not Hydnum fragile Pers. ex Fr., Syst. mycol. 1: 417. 1821 = Odontia macrodon (Pers. ex Fr.) Bourd. & Galz., cf. Lundell in Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. Fasc. 45-46: 1. 1954; not Hydnum fragile Petch in Ann. roy. bot. Gdns Peradeniya 7: 287. 1922.—Sarcodon fragilis (Fr.) P. Karst. mRev. mycol. 3 (No. 9) : 31 Jan.. 1, 1881 & in Medd. Soc. F. Fl. fenn. 6: 16. 1881. — Hydnum friabile Rostr. in Medd. For. Svampek. Fremme 2: 94. 1920 (avowed name change). — Type: (probably not in existence) represented by Fries, Sverig. ätl gift. Svamp. pl. 89. 1860 (selected).

Sarcodon reticulatus Banker in Mem. Torrey bot. Cl. 12: 139. 1906. — Hydnum reticulatum (Banker) Lloyd, Mycol. Writ. 4: Lett. 54, 7. 1915. — Type: not seen (NY).

Hydnum virginianum Murrill in Bull. Torrey bot. Cl. 67: 276. 1940. — Type: not seen (FLAS).

Descriptions.—Banker in Mem. Torrey bot. Cl. 12: 139. 1906 (Sarcodon reticulatus) ; Boudier, Icon, mycol. 4: 86. 1911 (Hydnum) ; Coker & Beers, Stip. Hydn. east. U.S. 34. 1951; Donk in Med. Nederl. mycol. Ver. 22: 63. 1933 (Sarcodon).

Illustrations.—Boudier, Icon, mycol. 1: pl. 168. 1904-1910 (Hydnum; very good) ; Bresadola, Icon, mycol. 21 ; pl. 1047. 1932 (Hydnum; mediocre) ; Coker in J. Mitch, sci. Soc. 41: pl. 54. 1926

(Hydnum; photogr., mediocre); Coker & Beers, Stip. Hydn. east. U.S. pl. 21.1951 (as preceding) ; FI. batava 19 : pl. 1470. 1893 (Hydnum fragile; unrecognisable, possibly some other species?) & 26: pl. 2028. 1924 (Hydnum infundibulum; very good); Fries, Icon. sel. Hym. 1: pl. 3 upper fig. 1867 (Hydnum; mediocre) ; Fries, Sverig. ätl. gift. Svamp. pl. 89. 1860 (Hydnum fragile; passable); Michael-Schulz, Führer Pilzfr. 3: pl. 305. 1927 (Hydnum infundibulum; fairly good) ; Rostrup in Medd. For. Svampek. Fremme 2: fig. 2. 1920 (Hydnum friabile; passable); Walty, Schweiz. Pilztaf. 3: pl. 65. 1947 (Sarcodon; colours misleading).

DIAGNOSTIC CHARACTERS.—Carpophores usually solitary. Pileus plano-convex, becoming depressed in centre,

margin incurved, finally undulate-lobate, surface felted-tomentose, even, often holding plant detritus, from

collapsing tomentum turning into glabrous, somewhat shining pellicle, long remaining pitted-tomentose to

finely fibrillose-scaly towards margin, pale alutaceous or flesh-coloured yellow-brown, fading out to whitish

near margin, finally dingy testaceous yellow-brown to darkish date brown. Stipe usually stocky, equal or

somewhat ventricose with tapering base, thinly felted, becoming fibrillose to squamulose, or glabrescent and

somewhat shining, pale dingy yellow-brown, at times suffused with some lilac tint, from base upwards turning

bistre with age, both fresh and dried with conspicuous whitish zone just below spines. Spines little or not

decurrent, pinkish when young (retaining this colour when dried), turning whitish, then some shade of grey.

Context thick, whitish in pileus, sometimes flushed with pinkish, pale yellowish brown with age, gradually

passing into pale bistre in stipe, not discolouring in KOH. Odour indistinct or agreeable when fresh, of

fenugreek when dried.

Habitat.—On acid sandy soil in pine woods, also in Cladonia-heaths. Distribution.—In most parts of the country, including the dunal region, but nowhere

common. Illustrative col lections.—Drente: Diever, Berkenheuvel, 6 X 1954,


Maas G. 10163 (L) ; Dwingelo, Schurenberg ven, 24 IX 1953, Miss M. H. Vaandrager (L). Exsiccati.—Ellis, North Amer. Fungi 929 (Hydnum fragile; L) ; Jaap, Fungi sel. exs. 142

(Hydnum; L) ; Litschauer & Lohwag, Fungi sel. exs. europ. 38 (Sarcodon; Herb. Donk, PR) ; Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. 2201 (Hydnum; C, PR, S) ; 2202 (Hydnum; C,

PR) ; Roumeguère, Fungi gall. exs. 2300 (or 2309?, illegible; Hydnum fragile; BR) ; Sydow,

Mycoth. germ. 1053 (Hydnum; K, L); Sydow, Mycoth. march. 1014 (Hydnum; K).

Bankera violascens (Alb. & Schw. ex Fr.) Pouz.

Hydnum violascens Alb. & Schw., Consp. Fung. 265. 1805; ex Fr., Syst. mycol. 1: 401. 1821; Epicr. Syst. mycol. 507. 1838; Hym. europ. 602. 1874. — Tyrodon violascens (Alb. & Schw. ex Fr.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 37: 91. 1882. —Sarcodon violascens (Alb. & Schw. ex Fr.) Quél. in C.R. Assoc. franç. Avanc. Sci. 11: 399 (13 of reprint). (1882) 1883; Ench. Fung. 189. 1886; Fl. mycol. 447. 1888.—Bankera violascens (Alb. & Schw. ex Fr.) Pouz. in Česka Mykol. 9: 96. 1955 & 10: 76. 1956. — Type: not known to be in existence. — Type locality: Germany, Oberlausitz, “Seer Busch; Dubrau; Hölle ; Arnsdorfer Berge” (Alb. & Schw., l.c. 265).

Hydnum cinereum Bull., Hist. Champ. France 1: 309. 1791 ; ex Fr., Syst. mycol. 1: 404. 1821 ; not Hydnum cinereum Bull, ex Fr. sensu Fr., Epicr. Syst. mycol. 508. 1838; Hym. europ. 604. 1874 = Phellodon confluens', nor Hydnum cinereum (Batsch) ex Pers., Mycol. europ. 2: 168. 1825 = Phellodon niger. — Hydnellum cinereum (Bull, ex Fr.) P. Karst, in Medd. Soc. F. Fl. fenn. 5: 41. 1879. — Calodon cinereus (Bull, ex Fr.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 37: 108. 1882. — Sarcodon cinereus (Bull, ex Fr.) Quél., Ench. Fung. 188. 1886. — Type: represented by Bull., Herb. France pl. 419. 1788.

Hydnum infundibulum Sw. in Svensk VetAkad. nya Handl. 31: 244. 1810; Fr., Obs. mycol. 1: 135. 1815; ex Fr., Syst. mycol. 1: 401. 1821; Elench. Fung. 1: 130. 1828; Epicr. Syst. mycol. 506. 1838; Hym. europ. 600. 1874. — Sarcodon infundibulum (Sw. ex Fr.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 37: 105. 1882. —Bankera infundibulum (Sw. ex Fr.) Pouz. in Česka Mykol. 9: 96. 1955 & 10: 76. 1956. — Type: probably non-existing. — Type locality: Sweden, “Seen in pine wood above Rotebro on the way to Upsala...” (Swartz, l.c. 245, translated).

Hydnum fusipes Pers., Mycol. europ. 2: 162. 1825. —Sarcodon fusipes (Pers.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 37: 105. 1882. — Sarcodon cinereus var. fusipes (Pers.) Quél., Ench. Fung. 189. 1886. — Type: “Hydnum fusipes. Mycol. Europ. Sect. 2. p. 162. Vogesia” [misit Mougeot] (L 910.263-922).

Hydnum torulosum Fr. in Öfvers. svensk VetAkad. Förh. 8: 53. 1851; Monogr. Hym. 2: 275. 1863; Icon. sel. Hym. 1: 5. 1867; Hym. europ. 600. 1874. — Sarcodon torulosus (Fr.) P. Karst, in Bidr. Känn. Finl. Nat. Folk 37: 106. 1882. — Type: (probably non-existing) represented by Fries, Icon. sel. Hym. 1: pl. 2 lower fig. 1867 (selected).

Hydnum ebneri Wettst. in S.B. kais. Akad. Wiss. math.-naturwiss. Cl. 94(1): 61. (1886) 1887. — Type: not known to be in existence. — Type locality: Austria, “Tirolia. In pinetis ad Siegmundslust prope Schwaz... ad Trins in valle Gschnitz...” (von Wettstein, l.c.).

Descriptions.—Albertini & Schweiniz, Consp. Fung. 265. 1805 (Hydnum) ; Konrad & Maublanc, Icon. sel. Fung. 5 : text to pl. 469. 1935 (Sarcodon) ; Persoon, Mycol. europ. 2: 162. 1825 (Hydnum fusipes).

Il lustrations.—Bresadola, Icon, mycol. 21: pl. 1043. (Hydnum infundibulum) ; pl. 1049 (Hydnum). 1932 (recognisable); Bulliard, Herb. France pl. 419 1788 (Hydnum cinereum; good) ; Fries, Icon. sel. Hym. 1 : pl. 2 lower fig. 1867 (Hydnum torulosum; very characteristic as to colour of context of stipe) ; Konrad & Maublanc, Icon. sel. Fung. 5: pl. 469. 1935 (Sarcodon; stipe and context probably too reddish) ; Nikolajeva in Not. syst. Sect. cryptog. Inst. bot. Komarovii Acad.

Sei. URSS 9: fig. 3. 1953 (Sarcodon; good); Persoon, Mycol. europ. 2 : pl. 20 fig. 4-6. 1825 (Hydnum fusipes, context too uniformly grey) ; Quélet in C. R. Assoc. franç. Avanc. Sci. 16: pl. 21 fig. 11. (1887) 1888 (Sarcodon; useless).


DIAGNOSTIC CHARACTERS.—Carpophores often with stipes connate at base or divided near top. Pileus convex,

becoming depressed in centre to umbilicate or infundibuliform, margin incurved, expanding with age, surface

thinly felted, even, not binding vegetable debris, from collapsing of tomentum turning into glabrous, somewhat

shining pellicle, subsequently rupturing radially, becoming scrupose to lamellate-scaly in centre, fibrillose-

scaly towards margin, sometimes also rupturing concentrically, giving rise to some degree of zonation, some

shade of pale lilaceous grey when young, passing into brownish flesh colour or pale purplish brown, with scales

often darker, margin concolorous. Stipe usually slender, tapering downwards, thin-felted, becoming fibrillose

to fibrillose-scaly, or glabrescent and somewhat shining, lilaceous whitish above, purplish brownish below,

fuscescent from base upwards with age except for narrow whitish zone just below spines. Spines decurrent,

white, then greyish. Context thick, white in pileus, flushed with lilac, purplish brown in stipe, discolouring

olive green in KOH. Odour indistinct when fresh, of fenugreek when dried.

Habitat.—Reported to grow in coniferous woods. Exsiccati.—Brinkmann, Westfäl. Pilze, Lief. 2: 100 (Hydnum; K, L; uncertain) ; Lundell &

Nannfeldt, Fungi exs. suec. praes. upsal. 353 (Hydnum; C,PR).

Both Donk (1933: 49) and Lundell (1947: 3) concur in considering Hydnum cinereum, as

redescribed by Fries in his ‘Hymenomycetes europaei’ (1874: 604), to be identical with Hydnum amicum (Phellodon confluens in the present paper). Hydnum cinereum Bull, ex Fr. (1821: 404), however, is something entirely different. Donk left the matter undecided as to what species Bulliard’s plate might be referred. Lundell, through an error, referred H. cinereum Bull, ex Fr. to Hydnum nigrum var. melilotinum at one time (1947: 1), to Hydnum amicum at another (1947: 3). Neither seems plausible to my mind, since both Phellodon niger and Ph. confluens have the stipe covered with a thick, soft tomentum. Bulliard’s plate 419, which in itself is beautifully executed, shows the specimens to have smooth stipes, at the most with a thin covering of superficial tomentum (“quelquefois aussi sa surface est pubescente...”) which once in a while may bind vegetable debris. The cut specimen also suggests that the context is homogeneous. This fact, coupled with the highly characteristic whitish zone at the top of the stipe depicted in a number of specimens, rather reminds of a Bankera. Further features, supporting the view that Bulliard’s plate actually represents B. violascens are (i) the clustered growth, (ii) the general colouring of the carpophores, (iii) the pronounced funnelled shape of the full-grown pileus in

some of the specimens, (iv) the radial striation, (v) the concentrical zones, and (vi) the rugosities in the centre of the pileus in others, (vii) the long stipe, which is unknown in Phellodon confluens, and (viii) the colour of its context, which excludes Phellodon niger. There is really only one point, viz. the spinous ‘head’ in the young stage which is rather a characteristic feature of species of the genus Phellodon, which may raise some doubt as to the identity of Bulliard’s fungus.

Bankera violascens is as yet unknown from this country, but may be expected to be found in the north eastern provinces.

Insufficiently known species

“HYDNUM VIOLACEU M” Hydnum violaceum Thore apud Pers., Traité Champ, comest. 249. 1818; ex Roques, Hist. Champ. 46. 1832. —

Sarcodon violaceus (Thore apud Pers. ex Roques) Quél. in C.R. Assoc, franç.

6o FUNGUS [Jaarg. 28 No. 1-4

Avanc. Sei. 22: 488 (5 of reprint). (1893) 1894. — Bankera violacea (Thore apud Pers. ex Roques) Pouz. in Česka Mykol. 9: 96. 1955 & 10: 76. 1956. — Type: not known to be in existence. — Type locality: France, Landes, near Dax, leg. J. Thore.

It is only because of Quélet’s words, “Spore... hyaline.” and, “Affine à violascens, A. et S. dont il n’est peut-être pas spécifiquement distinct.” that Hydnum violaceum is discussed under Bankera. But, while it is uncertain whether Quélet’s conception of the species squares with Thore’s Hydnum violaceum, it is impossible to ascertain from the scanty description whether the original species belongs to Bankera at all. The description of the context as “ . . . cassante,

tendre...” would point to the species being either a Bankera or a Sarcodon, the violaceous colours rather suggest S. commutatus Bourd. & Galz., but the description of the taste as very agreeable eliminates this thought.

REFERENCES

BLYTT, A. & ROSTRUP, E. 1905. Norges Hymenomyceter. In Vidensk. Selsk. Skr. 1. Math.- naturvidensk. Kl. 1904, No. 6.

DONK, M. A. 1933. Revision der niederländischen Homobasidiomycetae-Apyllophoraceae. Teil II. In Med. Nederl. mycol. Ver. 22.

-------- 1956. The generic names proposed for Hymenomycetes—V. “Hydnaceae”. In Taxon 5: 69-80, 95-115.

LUNDELL, S. 1947. Hydnum amicum Quél. In Lundell & Nannfeldt, Fungi exs. suec. praes. upsal. Fasc. 29-30: I. -------- 1947· Hydnum nigrum Fr. var. melilotinum (Quél.). In Lundell & Nannfeldt, Fungi exs.

suec. praes. upsal. Fasc. 29-30: 3. POUZAR, ZD. 1956. Prispěvek k poznani našich kloboukatých lošakŭ. In Česká Mykol. 10: 65-76. SNELL, W. H., DICK, E. A., JACKSON, H. A. C. & TAUSSIG, M. 1956. Notes on the pileate Hydnums. III. In Lloydia 19: 163-175.

INDEX

Bankera Coker & Beers ex Pouz. ……………………..56 Hydnellum amicum (Quél.) Ragab ……………….. 49 fuligineo-alba (Schmidt ex Fr.) Pouz 56,57 cinereum (Bull. ex Fr.) P.Karst …………………. 58 infundibulum (Sw. ex Fr.) Pouz. …………….. 58 cyathiforme (Schaeff. ex St.-Amans) P.Karst 54 violacea (Thore apud Pers. Ex Roques) Pouz 60 graveolens (Pers.) P.Karst …………………………… 51 violascens (Alb. & Schw. Ex Fr.) Pouz …………….. 58 melaleucum (Fr. ex Fr.) P.Karst ……………….….. 50 Calodon subgen. Phellodon (P.Karst.) P. Karst . 48 nigrum (Fr. ex Fr.) P.Karst ………………….………. 53 amicus (Quél.) Quél. ………………………………….. 49 velutinum var. scrobiculatum ………………………. 54 cinereus (Bull. ex Fr.) P.Karst ……………………… 58 var. zonatum 54,55 cyathiformis (Schaeff. ex St.-Amans) Quél. 54 Hydnum [sect.] Phellodon (P.Karst) J. Schroet. 48 var. candicans (Fr.) Quél. …………………………… 55 amicum Quél ……………………………………………... 49 var. variecolor (Secr.) Quél. …………………………. 55 candicans Fr. …………………………………………….. 55 graveolens (Pers.) Quél. ………………………………… 51 cinereum (Batsch.) ex Pers. …………………………... 53 f. nigricans Bourd. & Galz. ……………………………. 51 cinereum Bull. ex Fr. …………………………………… 58 f. ramosus Bourd. & Galz. ……………………………….51 cinereum Bull. ex Fr. Sensu Fr. 1838 ……………. 49 var. melaleucus (Fr. ex Fr.) Quél. …………………….50 confluens Peck …………………………………………… 49 melaleucus (Fr. ex Fr.) Quél. …………………………. 50 confluens Pers. …………………………………………… 49 niger (Fr. ex Fr.) Quél. ………………………………….. 53 coriaceo-membranaceum Schw. ………………….. 55 var. melilotinus (Quél.) Quél. ………………………… 53 cyathiforme Fr. ………………………………………….. 54 pullus (Schaeff. ex Pers.) Quél. ……………………… 50 cyathiforme Schaeff. ex St.-Amans ……………... 54 tomentosus (L. ex Fr.) R. Maire …………………….. 54 var. obscurum Sacc. …………………………………… 55 variecolor (Secr.) Quél. ………………………………… 55 ebneri Wettst. …………………………………………… 58 “variicolor” (Secr.) Quél. ………………………………. 55 fragile Fr. …………………………………………………. 57 fragile Pers. ex Fr. ……………………………………… 57

Nov. 1958] R. A. MAAS GEESTERANUS: Stipitate Hydnums—III 61 Hydnum fragile Petch 57 Phellodon amicus (Quél.) Banker . . 49 - friabile Rostr. 57 - brunneo-olivaceus Coker & Beers . 48 -fuligineo-album Schmidt ex Fr. . . . 56 - brunneoroseus Snell, Dick & Jackson 48 —fuscum foetens Secr. 51 — carnosus Banker 48 - fusipes Pers. 58 — cokeri Banker 49 — graveolens (Pers.) Fr. 50 - confluens (Pers.) Pouz. 48, 49 var. candicans (Fr.) Bres. 55 - coriaceo-membranaceus (Schw.) Banker 55 var. sub zonatum Peck 55 — cyathiformis (Schaeff. ex St.-Amans) P. Karst 54 subsp. melaleucum (Fr. ex Fr.) Légué 50 — infundibulum Sw. ex Fr. 58 - delicatus (Schw.) Banker . . . . 49 — leptopus Pers. 55 - ellisianus Banker 49 var. graveolens Pers. 50 — graveolens (Pers.) P. Karst 51 var. pullum (Schaeff.) ex Pers. . . 50 - melaleucus (Fr. ex Fr.) P. Karst. 48, 50 - melaleucum Fr. ex Fr. 50 f. candicans (Fr.) Nikolajeva . . . 55 — melilotinum Quél. 53 — f. graveolens (Pers.) Nikolajeva . . 51 — myriopedum Blytt 53 f. major Donk 51 - nigrum Fr. ex Fr. 48, 53 f. violascens Donk 51 var. melilotinum (Quél.) Lundell . 53 - niger (Fr. ex Fr.) P. Karst. 48, 53 var. olivaceo-cinereum Sacc. . . . 53 — f. melilotinus (Quél.) Donk . . . 53 subsp. *H. melilotinum (Quél.) Sacc. 53 - pullus (Schaeff. ex Pers.) Banker . . 50 — olivaceo-nigrum Secr. 53 — putidus (Atk.) Banker 48 -pullum (Schaeff. ex Pers.) Duby . . 5o, 53 - tomentosus (L. ex Fr.) Banker . . 49, 54 — pullum Sw. 50, 53 - vellereus (Peck) Banker 49 - reticulatum (Banker) Lloyd . . . . 57 Sarcodon amicus (Quél.) Quél. 49 suberosum var. cinerea Batsch . . . . 53 — cinereus (Bull, ex Fr.) Quél. 58 - “subtomentosum” L. ex Fr. 54 — var. fusipes (Pers.) Quél. 58 - sub zonatum (Peck) Lloyd 55 - commutatus Bourd. & Galz. . . . 6o — tomentosum L. ex Fr. 54 - fragilis (Fr.) P. Karst. 57 — var. atro-album Alb. & Schw. ex Pers. 50 - fuligineo-albus (Schmidt ex Fr.) Quél. 57 — torulosum Fr. 58 - fusipes (Pers.) P. Karst. 58 — variecolor Secr. 55 - infundibulum (Sw. ex Fr.) P. Karst. . 58 — vellereum Peck 49 - reticulatus Banker 57 — violaceum Thore apud Pers. ex Roques 59 torulosus (Fr.) P. Karst. 58 — violascens Alb. & Schw. ex Fr. . . . 58 - violaceus (Thore apud Pers. ex Roques) Quél. 59 — virginianum Murrill 57 — zonatum Batsch 55 -violascens (Alb. & Schw. ex Fr.) Quél. 58 — zonatum Batsch sensu Batsch 1789 55 vox. fuligineo-albus (Schmidt ex Fr.) Quél. 57 Odontia macrodon (Pers. ex Fr.) Bourd. & Galz. 57 Tyrodon fuligineo-albus (Schmidt ex Fr.) P.Karst. 56 Phellodon P. Karst 48 - alboniger (Peck) Banker 48 — violascens (Alb. & Schw. ex Fr.) P. Karst. 58

OBSERVATIONS ON GASTEROMYCETES—VII

Geastrum vulgatum Vitt.—A more definite name for Geastrum rufescens auct.

J. T. Palmer

The Hartley Botanical Laboratories, The University, Liverpool, England.

(With 1 Map, 2 Photographs and 14 Text-figures)

Vittadini’s material of G. vulgatum in Saccardo’s Herbarium at Padua is reexamined and a Lectotype is designated. A comparison is made with freshly collected G. vulgatum from Derbyshire, England, exsiccata in the Rijksherbarium, Leiden, in Herb. Kew and from Central Germany. A distribution map is given for Britain based on material in Herb. Kew. The literature is reviewed and the confusion with G. coronatum Pers., G. rufescens Pers. em. Kits van Wav. and G. triplex Jungh. is discussed. G. rufescens auct., G. schaefferi Vitt. and Plecostoma rufescens Desv. are considered synonyms.

Introduction.—Kits van Waveren (1926) was the first to demonstrate clearly that Geastrum rufescens Pers. is the valid name for the earthstar commonly known as G. fimbriatum Fr., although this had been suggested by earlier authors. The void created by this name change was filled by G. schaefferi Vitt., which appeared at the time to be the only available synonym.

Van Eyndhoven (1937a) endeavoured to locate authentic material of G. schaefferi but found that no specimens existed in the Vittadini Herbarium at Turin. He rightly commented: “ . . . der endgültige Beweis ist unmöglich, weil die Type von VITTADINI nicht mehr besteht.”

Discussing this problem in correspondence, Dr V. J. Staněk suggested that, from his perusal of the original diagnoses and figures (Vittadini 1842) G. vulgatum Vitt. made a better fit for G. rufescens auct. than did G. schaefferi. At the time, the author considered that the nomenclature of this well known fungus should not be further disturbed but, subsequently, he observed in Petri (1909) a reference to the Type specimen of G. vulgatum being extant in Saccardo’s Herbarium at Padua.

The Vittadini specimens were examined by the author and were found to match well material collected abundantly in Derbyshire, England, and determined as G. rufescens Pers. before he was aware of the true identity of this species. There would appear to be no material of G. schaefferi at Padua, and the writer has been unable to trace authentic specimens of either species elsewhere. The name G. vulgatum was used (Palmer 1955) when referring to this problem. The present paper is, therefore, intended to place the facts on record for the guidance of other workers.

Technique.—Micro-characters have been examined in Erythrosin Ammonia (Palmer 1955). All drawings are free-hand with components measured using an ocular micrometer. The

formulae show minimum, average and maximum measurements. Acknowledgments.—The author is grateful to Professor N. A. Burges, in whose Department

this work has been mainly carried out ; to Professor C. Capelleti, Director of the Herb. Micol. P.

A. Saccardo, Padua, for the loan of type

Nov. 1958] J. T. Palmer: Observations on Gasteromycetes—VII 63

and other material ; to Professor Beniamino Peyronel (Turin) for his views on the origin of the Padua specimens; for the loan of exsiccata to Dr R. A. Maas Geesteranus and the Director of the Rijksherbarium, Leiden, and, also, to Dr R. W. G. Dennis and the Director of the Royal Botanic Gardens, Kew; for material of G. vulgatum to Mr S. S. Bates (Stalybridge) and Dr H.-H. Handke

(Halle/Saale) ; to Dr V. J. Staněk (Prague) whose suggestion initiated this research and to Messrs. J. Bloor and C. E. Fenton for the photography. The writer is also indebted to Miss D. Buckley (Oldham) who originally informed him of the Derbyshire habitat.

VlTTADINl’S SPECIMENS

There are two sheets, both inscribed: “Geaster vulgatus Vittad. hab. a Vittadini. Mediol. 1841” and, also, but by a different hand in red ink, one is marked “TIPO” and the other “G. fimbriatus

forma”. There are no numbers or other means of reference.

Origin of specimens and script.—Photographs of the inscriptions were submitted to Professor Beniamino Peyronel, who informs the writer that “Mediol. 1841” would refer to the specimens being collected near Milan (Mediolanum) in 1841 whilst the writing in red ink probably belongs to Petri. With regard to the inscriptions “Geaster vulgatus etc.”, Professor Peyronel, after careful comparison with numerous samples of handwriting, considers that these, although by the same person, were not written by Vittadini but by Saccardo. This view is strengthened by the pencil notes and spore figures on the label of the syn-type, which corresponds well with Saccardo’s habits. He suggests that Saccardo, in his younger days, may have written to Vittadini for material of his new species and received the specimens of G. vul-

gatum accompanied by a letter, which were then carefully spread out on a sheet, noting the date and place of collection (Mediol. 1841) and emphasizing that they had been received from

Vittadini himself [hab(ui) a Vittadini].

Lectotype (Photo 1 and Figs. 1, 2 and 4).—The sheet marked “TIPO”, which is herewith

formally designated as the Lectotype, had two half specimens glued to it. The upper specimen shows external and the lower internal features but the latter has the upper part of the

Endoperidium and the Columella missing. Overall dimensions: 3.4-3.9 cm. Exoperidium with five remaining

(probably eight original) narrowly to broadly acuminate rays, divided for about two-thirds of the radius and

arched at the base. Mycelial Layer mainly formed of a thin, closely adhering weft of mycelium and soil particles,

partly missing to expose an ochraceous-red to almost apricot Fibrillose layer of a tough consistency, somewhat

stippled purplish-brown in parts on the upper surface where the remnants of the Fleshy layer have weathered

away. Endoperidium 1.2 x 1.2 cm., obturbinate, although possibly distorted by sectioning and pressing,

brownish-fawn, thin and papery, shortly stipitate, with what appeared to be a slight Apophysis. Mouth

indeterminate, edged with a fringe of fibrils. Pedicel 1.5 x 1.5 mm., spreading out above to form the basal part

of the Endoperidium and continuing upwards to form a short umbo within the Endoperidium, which would form

the base of the Columella, now missing. Gleba dark-brown, consisting of threads and the powdery spore mass.

Capillitium yellowish-brown, thick-walled- with the lumen almost completely obscured by the wall, sinuous and

irregular, occasionally encrusted, comprising simple threads with frequent short branches towards the tips,

64 FU NGUS [JAARG. 28 No. 1-4

Vittadini specimens in Herb. Micol. P. A. Saccardo: x—Lectotype; 2—Syn-type.

measuring 640-990-1280 μ long, tapering gradually to 1.5-2.2 μ diam, blunt tips at each end with a maximum

thickness of 5-7.3 μ. Spores brown, globose to subglobose, finely verrucose to rough, 1-guttulate, 3.2-3.9-4.1 x

3.1-3.6-3.8 μ, with verrucae 3.7-4.4-4.7 x 3.5-4.1-4.5 μ.

Syn-type (Photo 2 and Fig. 3).—This sheet bears a single half specimen glued so as to show

the external endoperidial characters.

Overall dimensions: 3.4 x 3.7 cm. Exoperidium with three (remaining) broadly acuminate rays, divided for

about two-thirds of the exoperidial radius, recurved and probably possessing an arched base. Mycelial Layer

mainly obscured but appeared to be formed of a persistent tfxin weft of hyphae and soil particles. Fibrillose

Layer tough and papery, light ochraceous-brown and mottled on the face where the remnants of the Fleshy

Layer have weathered away. Endoperidium 1.3 x 1 . 2 cm., sepia-brown, papery, shortly stipitate, rather badly

damaged. Mouth obscured but appeared to be indeterminate. Pedicel short, 1.5 x 1.5 mm. Gleba dark-brown,

formed of threads and the powdery spore mass. (The Columella could not be examined as that side of the

specimen was glued to the sheet.) Capillitium yellowish-brown, lumen only faintly apparent, often obscured,

rough or encrusted, formed of simple threads, 790-1010-1390 μ, gradually tapering to blunt ends, 1.9-2.8 μ

diam, at the tips, with a maximum thickness of 7-9.5 μ. Spores brown, globose to subglobose, 1-guttulate,

minutely verrucose, 3.2-3.8-4.1 x 3.1-3.7-4.2 μ, with verrucae 4.1-4.6-5.3 x 4.1-4.4-5.1 μ.


Observations.—Both specimens appeared to have been somewhat old when collected, judging by the mottled surfaces of the rays, which the author has frequently found in old, weathered peridia from Derbyshire and elsewhere. In all existing characters, the two Vittadini specimens made an excellent match with freshly collected Derbyshire material.

GEASTRUM V ULGATU M F ROM DERBY SHIRE , ENGLAND

In Britain, G. vulgatum seems to be far from common but the writer has collected it annually in October and November during the past five years on the lower parts of the scrub-covered side of Win Hill, near Ashopton, which is situated in the High Peak district of Derbyshire. The geasters develop in open grass patches surrounded by Pteridium aquilinum, particularly in exposed soil where the turf has been broken away, probably by grazing cattle scrambling up the hillside. Adjacent to these patches are clumps of Rubus and occasional individuals of Crataegus and, whilst strongly acid moorland conditions prevail towards the summit with a rich vegetation of Calluna vulgaris and Vaccinium myrtillus, more basic conditions are to be found where the geasters occur. This particular habitat faces due east and is rather exposed, whilst the lime-stone dales for which Derbyshire is noted occur about 10 km. to the south. Further specimens of G.

vulgatum have also been found on a grassy bank at Hathersedge in similar country about 5 km. to the south-east, whilst the author has examined specimens collected at Skircoat, Halifax, Yorkshire, some 40 km. to the north, which see under Herb. Kew.

The earthstars at Win Hill have been collected in nearly all stages of development: from unexpanded peridia to old, weathered individuals of the previous year or earlier. The author was particularly interested to ascertain the form of the Basidium in view of the structures which he has observed for G. rufescens and G. triplex (Palmer 1955). Unfortunately, whilst eggs ranging in size from a few millimetres to 2.3 cm. diam. were located, the glebae were found to be either too immature with only a few broadly clavate structures suggesting primordial basidia or in an advanced stage of deliquescence with no basidia remaining. The Tulasnes (1842) illustrated basidia of the more usual form for G. rufescens Pers. (very probably the species now under discussion) but, in view of the peculiar structures found for the two species mentioned, fresh observations are desirable.

Unexpanded peridia.—The eggs of G. vulgatum develop hypogeously in the clayey soil amongst grass roots with only a few thin, white strands of mycelium in evidence, although not infrequently adjacent to the decaying roots of trees whose upper parts have long disappeared. Rarely more than two or three eggs (or expanded peridia) are found adjacent to each other and, shortly before their expansion, the upper surfaces of the eggs can be seen projecting through the soil. The sparsity of the mycelium is in marked contrast to the abundant mycelial development in the habitats of G. rufescens and G. triplex, where individuals growing amongst leaf-mould in thickets often have an area of up to one square metre heavily impregnated with their conspicuous mycelium.


The eggs examined have varied up to 2 x 2.3 cm., depressed-globose in form, usually slightly concave at the

base. Mycelial Layer a thin weft of hyphae with adhering soil particles, usually only remaining as a few patches

where exposed, with the Fibrillose Layer of a striking flesh-pink colour. Gleba white but deliquescing before

expansion to form a dry, dark-brown mass of powdery Spores and Capillitium. During expansion, the Mycelial

Layer usually strips from the surface of the Fibrillose Layer and may be found partly lining the cavity which the

egg has previously occupied, although soon disappearing. The rays then recurve and the base becomes arched,

thus raising the fungus until it stands upon the flat surfaces of the Fleshy Layer, which often become curled

beneath.

FRESHLY-EXPANDED PERIDIA.—On completion of expansion, peridia measure from 3 to 8 cm. in

diameter of their extended rays, with a height of 2.5-4.5 cm., and are frequently considerably misshapen.

Exoperidium Assuring into 5-7-8 rays, extending inwards for about two thirds of the radius of the expanded

peridium, generally broadly acuminate but frequently with the main rays subdividing, which then result in

narrower structures. The base is usually prominently arched. Mycelial Layer thin, scurfy, admixed with a thin

layer of the clayey substratum, and usually stripping from the rays during expansion. Fibrillose Layer tough and

leathery, creamy-tan with a fleshy tint at first, varying to a deep flesh-pink which becomes ochraceous on

drying. Fleshy Layer at first pallid, becoming a deep flesh-pink, varying up to 4.5 mm. thick, finally much

cracked and fissured, giving the rays an irregular appearance, friable and soon flaking away. There was no

recognizable taste or odour. Endoperidium 7-14 x 8-20 mm., light buff to greyish-brown, varying from

subglobose to depressed-globose or urceolate, appearing to be sessile whilst fresh but, as the Fleshy Layer

shrinks or weathers away, becoming shortly pedicellate. Mouth indeterminate, concolorous with the

Endoperidium, fimbriate, plane to bluntly conical, finally becoming irregularly torn (‘toothed’ or ‘dentate’) and

gaping. Pedicel 1-2 mm. tall, formed from the Fibrillose Layer which thickens into a short, solid, cylindrical to

flattened structure, usually narrower in the middle, which broadens out above to form the shallow base of the

Endoperidium and also continues up within the Endoperidium to form the hard dome-like core, ca. 3 mm. diam.,

which is the foundation of the Columella. Gleba dark brown, formed of the Spore mass and Capillitium, with the

Columella broadly elliptical to globose, frequently reaching to just beneath the Mouth.

DRIED AND WEATHERED EXPANDED PERIDIA.—Their appearance may vary considerably but, in

the majority, the Fleshy Layer had disappeared or only remained on the surface of the tough Fibrillose Layer,

which was typically mottled with purplish or rufescent tinges, as a few dried-up fragments. However, if freshly

expanded specimens are carefully dried, the Fleshy Layer contracts to form a fairly uniform brown skin

although, even here, some cracking and flaking may occur. Such specimens readily soak out with the Fleshy

Layer regaining almost its original thickness.

In most instances, all traces of the Mycelial Layer had disappeared and the undersurface of the Fibrillose

Layer had weathered to a silvery white colour, not unlike that of the hygroscopic series, i.e. G. floriforme Vitt., G.

recolligens (Woodw. ex Sow.) Desv., etc. Old weathered specimens often have their ray surfaces green with

Algae, and the author has found several very old peridia, obviously from the previous year, which exhibited

strong hygroscopic tendencies with the rays inclosing over the Endoperidium on drying (Fig. 13) and expanding

when moistened (Fig. 14). The Endoperidium is shortly stipitate, although the Pedicel is usually rough and

shaggy with the desiccated remains of the Fleshy Layer. When dry, the Endoperidium is frequently found to have

contracted to expose a basal Apophysis (Fig. 12), which is rarely as prominent as with G.coronatum Pers. (= G.

limbatum Fr.). It seems to be due to the rigid nature of the basal part of the Endoperidium, where the upper part

of the Pedicel merges into and forms the endoperidial layer, and its degree of prominence depends upon the

thickness of this structure. MICROSCOPICAL CHARACTERS.—Whilst no actual basidia have yet been observed by


the author, he has found broadly clavate structures measuring 18 x 1ο μ in the hymenia of several very young

eggs, although they did not develop further. Capillitium yellowish-brown, with prominent to indistinct lumen,

nodular to rough, simple, with occasional short branches towards the tips, 600-1200-1900 μ long, tapering to

blunt ends, 1—2 μ diam., with a maximum thickness of 5.7-10 μ. Spores brown, globose, punctate to finely

echinulate, occasionally with what appeared to be a short apiculus, the contents indistinct but occasionally

appearing to be 1-guttulate, 3.1-3.8-4.5 x 3.1-3.8-4.5 µ, with verrucae 3.2-4·5-5·2 x 3.2-4·4-5·2 μ.

COLLECTIONS

Rijksherbarium, Leiden.—Four collections were examined and were found to be correctly

determined either as G. vulgatum or under a synonym:

1. H. L. B. Nederland No. 951.176-341. ’t Joppe bij Gorssel, Gelderland. 18.9.1951. Leg. W. J. Reuvecamp and det. G. schaefferi.

The single specimen, whilst with an immature gleba, was quite typical of G. vulgatum and had a strong

‘fruity’ smell.

2. H. L. B. Nederland No. 952.288-238. As above. Leg. 26.10.1952.

Similar in size and other characters to the Type and British material of G. vulgatum except for the more

pronounced Pedicel, with a decidedly ‘fruity’ odour. Spores brown, globose to occasionally ellipsoid, rough to

finely nodulose, 1-guttulate, 3.8-4.1-4.4 x 3.7-3.9-4.1 μ, with verrucae 4·3-4·6-5.2 x 4.1-4.4-4.8 μ.

3. H. L. Β. Nederland No. 958.021-439· Bergen, near café Duinvermaak, Noord-Holland. 28.10.1956. Leg. W. Prud’homme van Reine. Det. G. L. van Eyndhoven as G. vulgatum Vitt. ( = G. schaefferi).

A large specimen with 8 acuminate to wedge-shaped rays which were incurved when dry and expanded on

soaking and an arched base. The exoperidium measured 9.5 cm. when fully expanded and the fungus had a

‘fruity’ smell. Mycelial Layer debris-encrusted. Fibrillose Layer pink. Fleshy Layer coloured the water pink during

soaking. Pedicel short and thick. Endoperidium with a low Apophysis. Spores brown, globose to subglobose, finely

to bluntly warted, occasionally with a faintly apparent guttule, 3.6-4.0—4.7 x 3.1-3.8-4.4 μ, with verrucae 4.1-

4.9-5.8 x 3.8-4.7-5.6 μ.

4· H. L. B. Nederland No. 954.139-071. Wassenaar, Stoephout, Zuid-Holland, on leaf-litter under Pinus and Quercus, 2.10.1954. Leg. and det. R. A. Maas Geesteranus as G. schaefferi.

A large specimen measuring 11 cm. diam. when fully expanded with a ‘fruity’ smell. Mycelial Layer debris-

encrusted, reddish-cream. Fibrillose Layer pink. Fleshy Layer swelled during soaking and was much fissured. The

pigment of the Fleshy Layer coloured the water pink during soaking. Pedicel short, thick and somewhat

flattened. Spores brown, globose to ellipsoid, 1-guttulate, rough to shortly asperate with blunt warts, 3.4-4.0-

4.5 x 3.4-4.0-4·5 µ with verrucae 3.8-4.7-5.3 x 3.8-4.6-5.1 μ.

Hartley BOTANICAL LABORATORIES, LIVERPOOL.—One collection (Herb. Mycol Lpol. No. 843) comprising three fully expanded specimens collected on needle litter in a mixed wood, Grosse Probstei, near Naumburg (Saale), Central Germany, October, 1953. Leg. and det. H.-H. Handke as G. rufescens Pers.

Exoperidium 6-8 broadly acuminate rays, curling inwards and twisted when dry, splitting two-thirds to the

centre, with arched base, 9 cm. diam. of expanded rays of largest specimen, with a maximum height of 5.7 cm.

Mycelial Layer a thin felted layer mixed with decaying needles and other debris, missing in parts, particularly at

the base,


Known distribution of Geastrum vulgatum Vitt. for England and Wales based on material at Kew and collected by the author.

and exposing the Fibrillose Layer, which is tough and flesh-pink in colour. Fleshy Layer a thin, dried-up fawn-

brown layer, broken up into patches. Pedicel 2.5 x 4 mm., somewhat flattened, shaggy with remnants of Fleshy

Layer, concave in the middle and forming a low Apophysis to the Endoperidium, which is buffy-brown, 1.8 x 2.5-

3.5 x 2·2 cm., depressed-globose. Mouth indeterminate, fimbriate, torn and gaping. Gleba dark brown. Columella

bushy and globose with an ovate, whitish core. Capillitium yellowish-brown, rough and encrusted, lumen faintly

apparent, simple, 7.2-8.1 μ maximum thickness, tapering to 2.2 μ blunt tips. Spores brown, globose to ellipsoid,

finely to bluntly warted, contents obscured, 3.3-4.1-4.9 x 3.3-4.0-4.9 µ with verrucae 3·9-4.8-5·4 x 3.8-4·7-5·4

μ.

The collection was quite typical for G. vulgatum.

Herbarium, Kew.—A large number of specimens, variously determined as G. rufescens, G. fimbriatum, etc., have been examined. Some are without locality or data whilst many are old and often firmly glued to the sheets, which makes their identification difficult. The following

collections with data appear to belong to G. vulgatum:

I. Berkeley British Fungi 1836—1843. 275. Geaster rufescens P. minor. Two desiccated specimens glued to a sheet.

Spores with verrucae 3.6-4.8-5.7 x 3.3-4.5-5.0 μ.

Hollos (1904) listed this exsiccatum under G. rufescens Pers., the species under discussion.


2. Thringstone, Leicestershire. Leg. M. J. Berkeley and det. as (in pencil) G. rufescens P. minor.


3. King’s Cliffe, Northamptonshire. Leg. M. J. Berkeley. Two specimens glued to a sheet and labelled "Geastrum”.

Spores with verrucae 3.4-4.6-5.0 x 3.4-4.5 4.9 μ.

Berkeley & Broome (1848) referred to specimens of a small form being abundant at Thringstone, Leicestershire, a few years before, and to a single specimen occurring at Cliffe. Berkeley (1860) listed the species: “In pastures. Bardon Hills, Leicestershire.

Northamptonshire.”

4. Maidenhead, Berkshire. Leg. Rev. G. H. Sawyer.

Spores with verrucae 4.2-4.5-4.7 x 4.0-4.2-4.5 μ diam.

5. Wrekin Woods, Shropshire. Oct. 1869. (Ex Herb. W. G. Smith).

The specimen is glued to the sheet and hence difficult to examine but from its Spores (with verrucae 4.5-5.3-

6.0 x 4.5-5.1-5.9 µ) it appears to be G. vulgatum. It smelled faintly aromatic.

The occurrence of this species in “fine condition” in the woods around the Wrekin was

mentioned by Smith (1873).

6. Colwyn Bay, North Wales. 1880. (Ex Herb. M. C. Cooke) Det. G. fimbriatus Fr.

The Gleba is aborted but the general appearance of the fungus, which is glued to the sheet,

suggests that it is almost certainly G. vulgatum.

7. Lubbenham, near Ashton, Leicestershire. 1881. (Ex Herb. Myc. Berkeleyanum). Spores with verrucae 4.4-5.1-5.6 x 4.1-4.9-5.5 μ.

8. On the bare ground under Acer pseudoplatanus, Skircoat, Yorkshire. 27. Oct. 1907. (Ex Herb. C. G. Crossland).


This record was mentioned by Mason (1921).

9. Gower, South Wales. 1920. Leg. De G. Langlois. Det. as G. fimbriatus. Spores with verrucae 3.4-4.7-5.2 x 2.9-4.5-5.1 μ.

10. Canterbury, Kent. 28.10.1938. Leg. Mrs F. Hyland. Spores with verrucae 3·3-4·3-5.2 x 3.3-4.3-5.2 μ.

11. Bisley, Surrey. Leg. A. H. Childs. 16.9.1951. Det. as G. rufescens Pers.

With a note of having had a ‘strong fruity smell’, which still persisted. Spores with verrucae 3.2-4.4-5.1 x 3.2-

4.3-4.7 μ.

12. Cuffley Great Wood, Hatfield, Hertfordshire. Leg. P. H. Gregory. 20.9.1953.

Spores with verrucae 2.8-3·4-4·1 x 2.8-3.6-4.1 μ, but they were pale in colour and appeared not to be fully

formed.

It was otherwise typical G. vulgatum. H ERBARIUM SACCARDO , PADUA .—In addition to the Vittadini specimens, two collections

determined as G. vulgatum and one as G. rufescens (Pers.) Fr. were received on loan :

7 o FUNGUS [Jaarg. 28 No. 1-4

1. C. A. J. A. Oudemans, Fungi Neerlandici Exsiccati No. 244. Geaster vulgaris (altered to ‘... vulgatus’) Vitt. In dunis maritimis Walachriae. 1877.

A half specimen, with the Endoperidium lying loose, of typical G. triplex. Mycelial Layer free from debris,

purplish-brown in colour and longitudinally fissured. Mouth determinate, fimbriate and ragged. Fleshy Layer

mainly disappeared with the remnants forming a brown, honey-combed structure towards the ray tips. Spores

brown, globose to ellipsoid, verrucose, 3.1-3.8-4.4 x 3.1-3.6-4.1 μ, with verrucae 3.1-4.5-5.3 x 3.2-4.3-5.1 μ.

Hollós (1904) listed the exsiccatum as G. vulgaris under G. rufescens (= G vulgatum). Van Eyndhoven (1937b) under WALCHEREN considered Oudemans’ material so determined in Herb. Groningen to be G. triplex.

2. Geaster rufescens (Pers.) Fr. Petri Fl. it. crypt. Gast. p. 86, f. 65. Veneto Prov. di Treviso, nei prati aridi a Fadalto. 27. agosto 1915. Leg. R. Pampanini.

Three typical specimens of the true G. rufescens.

3. Geaster vulgatus Vittad. selve conifere del Trentino, with “G. fimbriatus forme” in red ink.

A single loose specimen typical of the true G. rufescens.

THE SPECIES IN LI TERATURE

Persoon (1801) included in his taxon G. rufescens two species but his exsiccata show that the species which he had in mind was the one subsequently called G. fimbriatum Fr. Fries (1829) appeared to be equally as vague in his diagnosis although, with his establishment of G. fimbriatum as distinct, he can be considered to have originated the tradition of G. rufescens for the present species. Berkeley & Broome (1848) stated that an authentic specimen from Persoon was identical with G. fimbriatum Fr. but, although subsequently referred to by several later authors, it was not until Kits van Waveren (1926) that a definite stand was taken and G. rufescens Pers. was re-established as the valid name for G. fimbriatum Fr.

G. vulgatum Vitt. and G. schaefferi Vitt. were published by Vittadini (1842) with G. vulgatum having priority of pagination. A comparison of the lengthy diagnoses seems to show little real difference between the two taxa except for the shape of the Columella, a character much used later by Petri (1909), and greater stress was laid on the reddish colouring of G. vulgatum.

However, G. schaefferi was later referred to G. rufescens by De Toni (1887), who considered G. vulgatum to be related to G. fimbriatum whilst resembling G. rufescens. Van Eeden (1874) recorded G. vulgatum for the Netherlands in pine woods at Bloemendaal. His description and figures suggest the true G. rufescens although his figs ƒ ƒ are peculiar stipitate forms. Van Eyndhoven (1937b) referred this record to G. rufescens, as he also does material from the same locality in Herb. Oudemans similarly determined. With the exception of Petri (1909), who listed

it for Lombardy (evidently the Vittadini specimens from the vicinity of Milan), G. vulgatum has been ignored by recent authors but G. schaefferi, as one of the few known synonyms of G. rufescens auct., was resurrected by Kits van Waveren (1926). However, the figures 52C, D and E for G. fimbriatum in Fischer (1933) are identical with G, H and I respectively of Vittadini (1842) Tab. I, Fig. I l l , Geaster vulgatus, Nob.

Several references have been made to G. rufescens var. minor Pers. which, in


Persoon (1801) as “colore dilutiore”, appears to be the small pale form of the true G. rufescens. As G. vulgatum has been generally considered to be a “large species”, small individuals, such as have been found by the author in Derbyshire, have been referred to this variety, see Berkeley British Fungi 275 under Herb. Kew. Smith (1908), giving “G. rufescens” as 4½ in. (11.5 cm.)

diam., stated the variety to be 5-8 segmented and less than half the size of the type. The genus Plecostoma was established by Desvaux (1809) for a section of Geastrum and

included the present species as P. rufescens. It has since fallen into oblivion.

CONF USI ON WI TH O THER GEASTERS

From the earliest times, G. vulgatum appears to have been confused with other species, not only the true G. rufescens but also with G. coronatum Pers. and G. triplex. This appears to be mainly due to the use of such inadequate criteria as “large size”, “rufescent colour”, "Endoperidium sessile or shortly stipitate” and “torn, toothed Mouth” instead of the more reliable debris-encrusted Mycelial Layer, indeterminate Mouth and the Spore characters nowadays considered to be of taxonomic importance.

Geastrum triplex Jungh.—G. vulgatum (as G. rufescens) has been considered to be a large species with a rufescent colour and a toothed Mouth but these characters are also possessed by other species, particularly G. triplex, which was first recorded for Britain by Smith (1873) as G. michelianus sp.n., where it is probably the commonest species, although Bryant (1782) gave good figures with the characteristic cup. G. triplex has a thick Fleshy Layer which fissures and flakes away, although not so readily as with G. vulgatum. The typical cup formed by the Fleshy Layer, absent in occasional specimens, should be considered an accidental feature and is generally missing from old, weathered individuals, which usually also have their Mouths torn and ragged. Van Eyndhoven (1937a) demonstrated that the Dutch specimens previously determined as G. rufescens or G. schaefferi were old peridia of G. triplex, although he later (1941) recorded it for the Netherlands. He also reported occasional shortly stipitate forms of G. triplex.

G. triplex develops epigeously as an onion-shaped egg and, at maturity, has a dark-brown, smooth Mycelial Layer free from debris which is typically longitudinally fissured. The Mouth is determinate although this character is not always clear in old specimens.

Geastrum rufescens Pers. em. Kits van Wav.—G. rufescens and G. vulgatum resemble each other macroscopically to some extent, although the former is typically saccate when expanded and rarely shows any sign of a pedicel whilst G. vulgatum usually has a prominently arched Exoperidium and a short Pedicel. Although the Fleshy Layer frequently has a reddish tinge, G. rufescens does not possess the flesh-pink colour so representative of G. vulgatum nor the inconspicuous Apophysis often found at the base of the Endoperidium. Further distinguishing characters of G. vulgatum would appear to be the typical reddish mottling on the upper surfaces of the rays of weathered specimens and the broadly elliptical to globose Columella whereas G. rufescens has a more slender structure. Microscopically, the two species can


usually be separated by the Spore dimensions with those of G. rufescens measuring 2.5-4.1 µ diam. with a prominent guttule, whilst G. vulgatum has Spores (2)-4-5-(6)μ diam. and the guttule, if present, is usually indistinct.

Geastrum coronatum Pers. (= G. limbatum Fr.).—The closeness of G. coronatum to G.

vulgatum was first pointed out by Scherffel (1896), who distinguished G. coronatum by its determinate Mouth and more strongly warted, darker Spores. Scherffel also considered G. vulgatum to have no Apophysis but Hollos (1904) and later authors recorded this structure for individuals and, of course, it is a criterion which is only apparent in desiccated specimens.

Lloyd (1902) considered G. vulgatum to be a reddish-brown fungus, stating that G. coronatum is usually a black species with a longer Pedicel and a differently shaped Endoperidium. The author has examined fresh material of G. coronatum which had a reddish coloured Fleshy Layer and, together with the Pedicel having been obscured by this structure, had been determined as G. triplex. G. coronatum is at first usually light in colour but tends to darken

with age.

TAXONOMI C CRI TERIA

Geastrum vulgatum can be recognised by (a) the depressed-globose shape of the egg ; (b) the arched form of the fully expanded Exoperidium; (c) the debris-encrusted Mycelial Layer, although this may be wholly or partly missing from expanded specimens; (d) the flesh-pink colour of the Fibrillose and Fleshy Layers; (e) the shortly pedicellate Endoperidium, which is immersed in the thick Fleshy Layer whilst fresh; (f) the indeterminate Mouth, usually becoming ragged; and (g) the finely warted Spores (3)-4-5-(6) μ diam. including the ornamentation.

Whilst disputed by some authors, desiccated individuals often have an Apophysis at the base of the Endoperidium. However, this structure must be considered to be more in the nature of an accidental feature in Geastrum, although very reliable for some species such as G. coronatum Pers. It disappears during soaking as the Endoperidium swells (see Figs. 12-14).

The flesh-pink colour of the Fibrillose and Fleshy Layers seems to be confined to G. vulgatum. Whilst other species of Geastrum often have a rufescent tinge to the Fleshy Layer, none have the delicate pink colour of G. vulgatum, which usually disappears with age and weathering. Large specimens examined by the author, and dried with the Fleshy Layer almost intact some years previously, were found to colour the water pink during soaking. A similar reaction was reported by Eberle (1956).

From the earliest days, opinion has vacillated between whether or not G. vulgatum (as G.

rufescens) had a sessile or a pedicellate Endoperidium. Whilst this variation of opinion may have had its origin in the confusion of the species with the true G. rufescens, it must be stressed that the Pedicel is not so prominent as is found with G. coronatum. The Fleshy Layer of G. vulgatum is exceptionally


thick and, consequently, it often does not weather away completely but may remain as dried-up ragged remnants, particularly around the Pedicel, which then assumes a rugged appearance. The majority of the pedicellate geasters appear to have sessile Endoperidia when freshly opened as their Pedicels are then obscured by the thick Fleshy Layers, and it is not until this layer has

partly or wholly disappeared that the typically stalked appearance is apparent. The “fruity odour” appears to have been first referred to by Coker & Couch (1928) who

remarked on the “very pleasant and decidedly fruity fragrance of the unopened and just opened plants.” Whilst no particular smell has been detected on the Derbyshire specimens, several exsiccata, particularly of the large forms, possessed a pleasant aroma. This may have been due to the thicker Fleshy Layer providing a more persistent habitat for the activity of microorganisms during desiccation, but another explanation must be sought for the odour of eggs and freshly expanded peridia of North American specimens.

The Columella was described by Vittadini (1842) as “longiuscula conica... crassa, ultra peridii centrum excurrens” and the author has found it to be a fairly robust, broadly elliptical to globose structure in those individuals which he has examined (Figs. 7-8). Petri (1909) reported this structure t o b e “longiuscula, conica” for G. vulgatum and “globosa, persistente” for G. rufescens (under which he placed G. schaefferi). He separated the latter on the Mouth being “fibroso-vel libroso-dentatum” but the torn or “dentate” Mouth, beloved by the old authors, is now realised to be an accidental trait of little significance.

Spore dimensions are usually given as 4-5 µ but Lloyd (1902) recorded 3-6 μ, Coker & Couch (1928) 3-4.4 µ and Smith (1951) 3-4.2 µ whilst the author’s measurements, including the ornamentation, are 3.7-4.4 μ for the Type material, 3.2-5.2 µ for the Derbyshire collections and 3.2-6.0 µ for specimens from elsewhere. Most authors have considered the Spores to be agutullate but the author has observed guttules in occasional Spores in most collections, although they are not so clear as in other species.

The Capillitium is generally given as being “thicker than the Spores”, with dimensions between 6 µ and 10 μ, although Coker & Couch (1928) gave 3.5-4.8 μ and Smith (1951) 3.5-6 µ. The author found the maximum thickness of the Capillitium to range from 5 µ to 9.5 µ for the Vittadini specimens and 5.7 µ to 10 μ for elsewhere.

Vittadini (1842) reported G. vulgatum: “In nemore della Rossa supra Roncaro, in ticinensi provincia, obvius. Autumno.” and G. schaefferi: “In nemore della Rossa, prope Roncaro circa Paviam. Autumno.” As was pointed out by Dr M. A. Donk, “ticinensi” is derived from “Ticinum”, the ancient name for Pavia (= Padua) and does not refer to the Swiss Canton Ticino, as indicated

by van Eeden (1874). Most European authors report the species from coniferous woods but British writers, starting with Berkeley (1860), have recorded it from pastures and wooded localities whilst Ramsbottom (1953) mentioned oak woods. Van Eyndhoven (1941) found G. vulgatum beneath Fagus sylvatica and Tilia, and the Skircoat collection recorded by Mason (1921) was under Acer pseudoplatanus. In North America, Coker & Couch (1928) reported G. vulgatum at the bases of oak stumps and Smith (1951) in debris on light soil around hardwood stumps. The author’s collections in Derbyshire were on rough pasture. The species does not appear to be known with certainty outside Europe and North America.

74 Fungus [Jaarg. 28 No. 1-4


CONCLUSI ONS

The discovery of authentic specimens of Geastrum vulgatum Vitt. has resulted in this species, whose taxonomic position was previously dubious, being definitely identified with G. rufescens auct. G. schaefferi Vitt., although similarly doubtful, had been considered a synonym of G. rufescens and was therefore promoted to fill the vacancy left when G. rufescens Pers. em. Kits van Wav. was established to be a different fungus. The absence of authentic specimens for G. schaefferi makes the true identity of this species uncertain, hence, as both taxa are of equal priority, the author considers that G. vulgatum Vitt. should be regarded as the valid name for G. rufescens auct. with G. schaefferi as a probable synonym.

Vittadini’s specimens of G. vulgatum are small forms similar to those found by the author in exposed grassland and differ markedly in size from peridia of G. rufescens auct., traditionally considered a large species, found in coniferous woods and similar habitats. A careful comparison between the grassland forms and those collected in woods shows no real difference. The development of smaller fruit-bodies in open areas should be attributed to the more luxuriant conditions prevailing in the rich humus of wooded areas and, even here, there was some overlapping of the gross dimensions of individuals examined from the two types of habitat.

Small, somewhat atypical, weathered forms of G. vulgatum with strong hygroscopic tendencies found by the author in Derbyshire (Figs. 13-14) emphasize the caution which should

be exercised when naming old peridia. And indeed, it is often as old battered sporophores, which would be discarded as being long past identification in other groups of the Higher Fungi, that geasters are frequently found.

REFERENCES

BERKELEY, M.J. 1860. Outlines of British Fungology. London. ------------ & BROOME, C. E. 1848. Notices of British Fungi. No. 378. Geaster rufescens P. In Ann. Mag. nat. Hist.,

Ser. 2, 2: 268. BRYANT, G. 1782. An historical account of two species of Lycoperdon. London. (Privately published) . COKER, W. C., & COUCH, J. N. 1928. The Gasteromycetes of the Eastern United States & Canada. Chapel Hill. DESVAUX, N. A. 1809. Observations sur quelles genres à établir dans la famille des Champignons. In Desv. J. Bot.

2: 88-105.

Geastrum vulgatum Vitt.—Macroscopic 1 x ; microscopic 1000 x. Figs. 1—4. Vittadini’s specimens: 1— ½ specimen of Lectotype showing internal characters; 2— ½ specimen of

Lectotype showing external characters; 3—Syn-type; 4—Spores and Capillitium of Lectotype. Figs. 5-14. Derbyshire material: 5—Spores and Capillitium; 6—External appearance of an egg; 7—Section of an

egg; 8—Section of a freshly expanded peridium; 9—Freshly expanded peridium with revolute rays and a plane, torn Mouth; 10—Freshly expanded peridium with a papillate Mouth; 11—Freshly expanded peridium with a much cracked Fleshy Layer and a torn Mouth; 12—Old expanded peridium with adhering desiccated remnants of the Fleshy Layer and a pedicellate Endoperidium with a slight Apophysis and a torn Mouth; 13—Old, weathered peridium with rays tightly inclosed and showing a contracted Endoperidium and an Apophysis; 14—The same specimen after soaking with the rays recurving and the Endoperidium “plumped out”.

76 FU N G US [JAARG. 28 No. 1-4

DE TONI, G. B. 1887. Revisio monographica generis Geasteris Mich, e tribu Gasteromycetum. In Rev. myc. 9: 61-77 & 125-133.

EBERLE, G. 1956. Erdstern - Nachlese. In Jahrb. nassau. Ver. Naturk. 92: 30-36. EEDEN, F. W. VAN, 1874. In Flora batava 15: Pl. 1165. EYNDHOVEN, G. L. VAN, 1937a. Geastrologische Notizen. 3. Die Frage des Vorkommens von Geastrum Schaefferi

Vitt, in den Niederlanden. In Med. Ned. myc. Ver. 24: 14-19. --------- 1937b. Uebersicht über die Verbreitung der Genera Geastrum, Myriostoma und Astraeus

in den Niederlanden. In Med. Ned. myc. Ver. 24: 20-48. ---------- 1941. Geastrum Schaefferi Vitt, in den Niederlanden gefunden. In Ned. kruidk. Arch. 51 :

380-384. FISCHER, E. 1933. In Engler & Prantl, Die Natürlichen Pflanzenfamilien, 2 Aufl., 7a. Leipzig. FRIES, E. M. 1829. Systema mycologicum, 3. Gryphiswaldae. PIOLLÓS, L. 1904. Die Gasteromyceten Ungarns. Leipzig. KITS VAN WAVEREN, E. 1926. De Nederlandsche Soorten der Genera Geaster, Myriostoma en Astraeus. In Med. Ned.

myc. Ver. 15: 85-128. LLOYD, C. G. 1902. The Geastrae. Bull. Lloyd Lib. 5, Myc. Ser. No. 2. 1-44. MASON, F. A. 1921. Geaster rufescens Pers. in Yorkshire. In Naturalist, Lond. 74-75.

PALMER, J. T. 1955. Observations on Gasteromycetes. 1-3. In Trans. Brit. myc. Soc. 38: 317-334· PERSOON, C. H. 1801. Synopsis methodica fungorum. Göttingen. PETRI, L. 1909. Gasterales. In Flora italica cryptogama. Pars I. Fungi. RAMSBOTTOM, J. 1953. Mushrooms & Toadstools. London. SCHERFFEL, A. 1896. Bemerkungen über Geaster-Arten. In Ber. deutsch, bot. Ges. 14: 312-323. SMITH, A. H. 1951. Puffballs and their allies in Michigan. Ann Arbor. SMITH, W. G. 1873. Starry Puff-balls. IV & V. In Gard. Chron. 577 & 608. --------- 1908. Synopsis of the British Basidiomycetes. London. TULASNE, L. R. & C. 1842. Sur les genres Polysaccum et Geaster. In Ann. Sci. nat. Bot., ser. 2, 18:129-141. VITTADINI, C. 1842. Monographia Lycoperdineorum. Turin. [Also (1843) in Mem. Ac. Torino, Ser. 2, 5: 145-237.]

Nov. 1958]

BOEKBESPREKINGEN

77

F. H. Möller, Fungi of the Faeroes

Part I, 1945, Basidiomycetes, 296 pp. + 3 coloured plates and map, Dan. Cr. 20— Part II, 1958, Myxomycetes, Archimycetes, Phycomycetes, and Fungi Imperfecti, with an Appendix to Part

1, 288 pp. + coloured plate, Dan. Cr. 60 — (Ejnar Munksgaard, Copenhagen).

Halverwege tussen Schotland en IJsland rijzen de Faröer-eilanden, kaal en boomloos, uit de zee op. Het klimaat is er erg vochtig. De jaarlijkse regenval bedraagt bijna 1500 mm en het regent er 3 van de 4 dagen. De gemiddelde maandtemperaturen lopen er weinig uiteen. Voor de koudste maand van het jaar, maart, bedraagt deze 3,1 °C en voor de warmste, juli, 10,8 °C.

Omtrent de fungi van deze tamelijk onherbergzame eilanden was vóór 1938 nog maar weinig bekend. In dat jaar is echter F. H. Möller in staat gesteld gedurende drie maanden (van half juni tot half september) er de mycologische flora te bestuderen. De resultaten van Möller’s onderzoek

zijn in een tweetal forse delen neergelegd. Het eerste deel dateert van 1945, terwijl het tweede pas in 1958 is verschenen. Daar beide delen samen een gesloten geheel vormen en ook het eerste deel nog in de boekhandel verkrijgbaar is, lijkt het me nuttig aan een bespreking van het laatste deel een overzicht van het eerste te laten voorafgaan, te meer daar hiervan nimmer een bespreking in Fungus heeft plaats gevonden.

Het meest belangwekkend uit sociologisch oogpunt zijn die soorten op de Faröer welke er op de natuurlijke weiden en de veenachtige terreinen voorkomen. Hieronder tellen we vele Clavaria’s, Hygrophori en rosesporigen. Andere hebben deze eilanden door toedoen van de mens bereikt, zoals de verstekelingen die met geïmporteerde kruidachtige planten, struiken of bomen of zelfs met timmerhout zijn meegekomen en voorts de coprophile soorten, gebonden aan de mest van geïmporteerde schapen en koeien. Het is merkwaardig dat de mestzwam Anellaria separata (= Panaeolus separatus = Panaeolus semiovatus) er de meest algemene hogere zwam is geworden ; ze komt er zelfs voor op de met stroplaggen bedekte daken van de huizen. Indien een bewoner van de Faröer dan ook wil zeggen dat iets erg algemeen is, bezigt hij vaak de uitdrukking: het is zo gewoon als „hunderland” op het dak. „Hunderland” betekent zoiets als hondenurine. Allengs is deze term, die van weinig appreciatie getuigt, zelfs synonym geworden met het Nederlandse „paddestoel”.

Het eerste deel bevat in totaal de uitvoerige beschrijving van 202 Basidiomyceten, verdeeld over 57 Ustilaginales en Uredinales, 22 Aphyllophorales (waaronder ook de 5 Boleten zijn gerangschikt) en 123 Agaricales met hun variëteiten en vormen. De meeste beschrijvingen zijn door een afbeelding begeleid. Verscheidene nieuwe soorten en variëteiten zijn opgenomen en het is te verwachten dat een deel daarvan op onze kale duinterreinen e.d. kan worden teruggevonden. In het supplement van deel twee worden nog eens 39 soorten toegevoegd aan groepen in het eerste deel behandeld.

Het tweede deel vangt aan met opmerkingen omtrent het verzamelen en met een uitvoerig overzicht omtrent de oecologie van de daarin behandelde species. De Discomyceten zijn grotendeels door Nannfeldt bewerkt, de Pyrenomyceten door Munk en de overige micromyceten door Buchwald. Exclusief het supple-


ment bevat dit deel de beschrijvingen van 311 soorten, waaronder natuurlijk ook weer vele „novae species” en meerdere nieuwe combinaties. De hoofdschotel wordt gevormd door 151 Ascomyceten en 140 Fungi Imperfecti. Een gekleurde plaat is in hoofdzaak aan kleine Discomyceten gewijd. Beide delen bezitten overigens een 300 figuren en foto’s.

Het werk is op zeer mooi papier uitgegeven. De afgebeelde soorten zijn goed herkenbaar. Volgens de geijkte terminologie: ongetwijfeld een werk waarvan het bezit zijn nut kan afwerpen. Dank zij Möller’s werk is het me trouwens reeds meermalen gelukt moeilijke gevallen tot een

oplossing te brengen.

H. S. C. Huijsman

Dr H. C. BELS-KONING en Drs P. J. BELS, Handleiding voor de Champignon- cultuur

295 pp., 94 fig. en foto’s, 29 tabellen, 1958 (Proefstation voor de Champignoncultuur, Horst, Limburg)

Het nog maar kort bestaande Proefstation voor de Champignoncultuur te Horst is wel zeer actief. Het heeft reeds het regelmatig verschijnende tijdschrift „De Champignoncultuur” opgericht en deed nu een Nederlands handboek voor de champignoncultuur het licht zien.

Na de laatste wereldoorlog is de champignoncultuur in Nederland snel in betekenis toegenomen. Er zijn nu reeds 650 kwekerijen die in 1957 gezamenlijk 1,6 miljoen kilo champignons op de markt brachten, waarvan 1 miljoen kilo werd uitgevoerd. Een goede voorlichting is bij deze moeilijke cultuur dus zelfs van algemeen belang.

Het boek dat keurig is uitgevoerd en vele goede foto’s, tekeningen en tabellen bevat, behandelt zeer vele facetten van de champignoncultuur : van selectie tot kostprijsberekening, van nomenclatuur tot kwekerijbouw. Vooral de lange tabellen met afwijkingen en ziekten en hun oorzaken of veroorzakers en de uitgebreide literatuurlijst zullen voor de kweker van groot nut zijn.

Enige opmerkingen van ondergeschikt belang zijn de volgende. Een index had in dit boek eigenlijk niet mogen ontbreken. Jammer dat niet duidelijk naar voren komt dat Agaricus de juiste genus-naam voor de champignon is en niet Psalliota. Het valt zeer te betwijfelen of

Clitocybe dealbata en Pleurotus mutilus (p. 186) identiek zijn. Overigens een boek om de auteurs en het Proefstation voor de Champignoncultuur geluk mee

te wensen.

C.BAS

I N HO U D

J. A. von Arx: Die systematische Stellung der Ascomycetengattung Armatella Theiss. et Syd. ……………………………….1

—-----------: Ueber den Ascomyceten Rhagadolobium cucurbitarum (Rehm) Theiss. et Syd ……………………………… 4

M. A. DONK: The generic names proposed for Hymenomycetes—IX. “Meruliaceae” and Cantharellus s.str …………. 7

--------- : Notes on resupinate Hymenomycetes—V ....................................................................................... ……16

J. GREMMEN: Taxonomical notes on mollisiaceous fungi—VI. The genus Pyrenopeziza Fuck. ……………………… ………37

H. S. C. HUIJSMAN: Note complémentaire à propos de Cystoderma superbum Huijsm ..................................... …..47

R. A. MAAS GEESTERANUS: The stipitate Hydnums of the Netherlands—III. Phellodon P. Karst, and Bankera Coker &

Beers ex Pouz ………………………………………………………………………………………………………………………………………… 48 J. T. PALMER: Observations on Gasteromycetes—VII. Geastrum vulgatum Vitt. — A more definite name for

Geastrum rufescens auct ……………………………………………………………………………………………………………….. 62

Boekbespreking ……………………………………………………………………………………………………………………………… 77

N ED ER LA N D S E M Y C O LO G I S C H E VER E N I G I N G

G. L. VAN EYNDHOVEN, voorzitter, Floraplein 9, Haarlem. Dr G. A. DE VRIES, onder-voorzitter, Beukenlaan 8, Baarn.

Mej. Dr A. JAARSVELD, secretaresse, Hugo de Vries-laboratorium, Plantage Middenlaan 2a Amsterdam-G. Mej. J. P. S. SMIT, penningmeesteresse, Nic. Maesstraat 135, Amsterdam-Z.

Giro Nederl. Mycol. Ver., Amsterdam, No 90902.

Dr M. A. DONK, Prins Mauritslaan 18, ’s-Gravenhage.

Dr R. A. MAAS GEESTERANUS, bibliothecaris, Rijksherbarium, Nonnensteeg 1, Leiden.

Still available:

A. F. M. Reijnders: Recherches sur le développement des

carpophores dans les Agaricales

1952. Amsterdam. Price Hfl. 10.— (Hfl. 2.50 for members)

Orders to be addressed to: Dr R. A. Maas Geesteranus, Rijksherbarium, Nonnensteeg 1, Leiden.

D R U K : H . V E E N M A N E N Z O N E N , W A G E N I N G E N

28e Jaargang No 1 -4 27 november 1958 FUNGUS · oder Schiffnerula v. Höhn, besitzen bitunicate...

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Transcript of 28e Jaargang No 1 -4 27 november 1958 FUNGUS · oder Schiffnerula v. Höhn, besitzen bitunicate...