Typification of some names of rust fungi described by A. P. de Candolle

Abstract

A. P. de Candolle (1778–1841) described numerous species names of rust fungi of the genera Aecidium, Puccinia and Uredo.
Original collections deposited in de Candolle’s herbarium (G) under the names concerned were examined in order to clarify
their typifications. Some types were classified as holotypes, according to Art. 9.1, Note 1. Other names are lectotypified or,
when necessary, neotypes are designated. The nomenclature of the names involved was disentangled, including clarification
of the correct current names, based on the currently valid Code (ICNafp). The new combination Uromyces bifrons is
introduced.

Full text

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Schlechtendalia 39, 2022
Typification of some names of rust fungi described by A. P. de Candolle
Uwe BRAUN
Abstract: Braun, U. 2022: Typification of some rust names described by A. P. de Candolle. Schlechtendalia 39: 54–63.
A. P. de Candolle (1778–1841) described numerous species names of rust fungi of the genera Aecidium, Puccinia and Uredo.
Original collections deposited in de Candolle’s herbarium (G) under the names concerned were examined in order to clarify
their typifications. Some types were classified as holotypes, according to Art. 9.1, Note 1. Other names are lectotypified or,
when necessary, neotypes are designated. The nomenclature of the names involved was disentangled, including clarification
of the correct current names, based on the currently valid Code (ICNafp). The new combination Uromyces bifrons is
introduced.
Zusammenfassung: Braun, U. 2022: Typisierung einiger von A. P. de Candolle beschriebener Rostpilznamen.
Schlechtendalia 39: 54–63.
A. P. de Candolle (1778–1841) hat zahlreiche Artnamen von Rostpilzen der Gattungen Aecidium, Puccinia und Uredo
beschrieben. In de Candolles Herbarium (G) unten den entsprechenden Namen deponiertes Originalmaterial wurde
untersucht, um deren Typisierungen zu klären. Einige Typen konnten, sprechend Art. 9.1, Note 1, als Holotypen eingestuft
werden. Andere Namen werden lectotypisiert oder es werden, wenn notwendig, Neotypen eingeführt. Die Nomenklatur der
involvierten Namen wird klargestellt, einschließlich Klärung der gegenwärtig gültigen Namen, entsprechend dem heutigen
Code (ICNafp). Die neue Kombination Uromyces bifrons wird eingeführt.
Key words: Aecidium, Puccinia, Uredo, herbarium G, typus, species.
Published online 29 Mar. 2022
Introduction
De Candolle (1805, 1815) described numerous rust fungi under the genus names Aecidium, Puccinia
and Uredo. The typification of most of his rust names has not yet been clarified. In the course of
nomenclatural studies on rust names, in connection with phylogenetic investigations, numerous
specimens deposited under the names concerned in the herbarium of A. P. de Candolle at G have been
examined. A few holotypes have been identified, and, in other cases, lecto- and neotypes are
designated. The nomenclature of the rust names involved is treated in detail. Some additional names
within the synonymy of the treated names are also lectotypified.
List of typified names
(1) Aecidium bifrons DC. [α. aconiti-lycoctoni], in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 246,
1805.
Holotype: France, “fl. fr. 662”, on Aconitum lycoctonum, [M. Chaillet], ex herb. de Candolle
(G00262469).
 Nigredo bifrons (DC.) Arthur, N. Amer. Fl. 7(11): 760, 1926.
 Uromyces bifrons (DC.) U. Braun, comb. nov. (Basionym: Aecidium bifrons DC., in Lamarck & de
Candolle, Fl. franç., Edn 3, 2: 246, 1805). MycoBank, MB833525.
 Uromyces aconiti-lycoctoni (DC.) G. Winter, Rabenh. Krypt.-Fl., Edn 2, 1.1: 153, [1884] 1881.
 Uromycopsis aconiti-lycotoni (DC.) Arthur, Résult. Sci. Congr. Bot. Wien 1905: 345, 1906.
= Uredo lycoctoni Kalchbr., Mathem. Természettud. Közlem. 3: 306, 1865.
 Aecidium lycoctoni (Kalchbr.) Sacc. & D. Sacc., Syll. fung. 17: 358, 1905.
 Uromyces lycoctoni (Kalchbr.) Trotter, Fl. ital. crypt., Fungi 4: 64, 1908.
= Uromyces aconiti Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 61 [1869-1870] 1870.
Notes: De Candolle, in Lamarck & de Candolle (1805), described Aecidium bifrons with α. aconiti-
lycoctoni and “. Aecidium ranunculi acris Pers. 210?”, which refers to Persoon (1801) and not to the
original description of Aecidium ranunculi-acris Pers. However, the name Aecidium bifrons is
nevertheless legitimate and not superfluous since Persoon’s name was only cited with question mark,
i.e., this name was not included in the original description. Therefore, only “α. aconiti-lycoctoni”
pertains to the protologue, with Aconitum lycoctonum as type host. There is only a single collection in
de Candolle’s herbarium which is in accordance with the protologue, so that this specimen can be

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classified as holotype (according to Art. 9.1, Note 1). Different names have been applied to the
common Aconitum lycoctonum rust, but Aecidium bifrons is the oldest valid name for this species.
(2) Aecidium geranii DC. [. Geranii rotundifolii], in Lamarck & de Candolle, Fl. franç., Edn 3, 2:
246, 1805.
Lectotype (designated here, MycoBank, MBT10006424): France, on Geranium rotundifolium, ex
herb. de Candolle (G00262492).
 Caeoma geraniatum Link, in Willdenow, Sp. pl., Edn 4, 6(2): 57, 1825, nom. nov. [non Caeoma
geranii (DC.) Schltd. 1824].
= Uredo geranii DC., in Lamarck & de Candolle, Syn. pl. Fl. Gall.: 47, 1806.
 Caeoma geranii (DC.) Schltdl., Fl. berol. 2: 128, 1824.
 Uromyces geranii (DC.) Lév., Ann. Sci. Nat., Bot., Sér. 3, 8: 371, 1847, nom. inval. (Art. 35.2).
 Uromyces geranii (DC.) Fr., Summa veg. Scand., Sectio Post.: 514, 1849.
 Coeomurus geranii (DC.) Kuntze [as “Caeomurus”], Revis. gen. pl. 3(3): 450, 1898.
 Nigredo geranii (DC.) Arthur, N. Amer. Fl. 7(11): 765, 1926.
= Trichobasis geranii Berk., Outl. Brit. Fung.: 333, 1860.
Notes: De Candolle (1805) described Aecidium geranii with α. Geranii pusilli and . Geranii
rotundifolii. The syntypes are preserved in de Candolle’s herbarium (α. Geranii pusilli, G00262492).
However, I prefer to designate de Candolle’s well-preserved material on Geranium rotundifolium as
lectotype.
(3) Puccinia laburni DC., in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 224, 1805.
Neotype (designated here, MycoBank, MBT10006425): France, on Laburnum anagyroides, 1821, A.
P. de Candolle, ex herb. de Candolle (G00262478).
 Uredo laburni (DC.) DC., Encycl. Méth. Bot. 8: 222, 1808.
 Uromyces laburni (DC.) G.H. Otth, Mitt. Naturf. Ges. Bern 531-552: 87, 1864.
Notes: Type material of Puccinia laburni collected before 1805 is not preserved. Therefore, a later
collection deposited in de Candolle’s herbarium is designated as neotype.
(4) Puccinia phyteumatum DC. [as “phyteumarum”, α. phyteumatis-spicati], in Lamarck & de
Candolle, Fl. franç., Edn 3, 2: 225, 1805.
Lectotype (designated here, Mycobank, MBT10006426): France, on Phyteuma spicatum, “fl. fr. 602”,
ex herb. de Candolle (G00262480).
 Uredo phyteumatum (DC.) DC., Encycl. Méth. Bot. 8: 222, 1808.
 Caeoma phyteumatum (DC.) Schltdl. [as “Caeoma phyteumatis”], Linnaea 1: 611, 1826.
 Uromyces phyteumatum (DC.) Niessl, Verh. Nat. Ver. Brünn 3: 114, 1864.
 Coeomurus phyteumatum (DC.) Kuntze [as “Caeomurus”], Revis. gen. pl. 3(3): 450, 1898.
 Telospora phyteumatum (DC.) Arthur, Résult. Sci. Congr. Bot. Wien 1905: 346, 1906.
Notes: De Candolle (1805) based the name Puccinia phyteumatum on collections on two host species,
viz., α. phyteumatis-spicati and . phyteumatis-orbiculare. Hence, it is necessary to designate a
lectotype for the species name. G00262480 is the only collection in de Candolle’s herbarium that
coincides with the protologue of α. phyteumatis-spicati. Therefore, this specimen is designated as
lectotype.
(5) Puccinia phyteumatum DC. . phyteumatis-orbiculare DC., in Lamarck & de Candolle, Fl. franç.,
Edn 3, 2: 225, 1805.
Holotype: France, on Phyteuma orbiculare, ex herb. de Candolle (G00262482).
Notes: The taxonomic rank of α. and . was not indicated in the original description. The cited
collection is the only specimen in de Candolle’s herbarium that is in agreement with the protologue.
Therefore, it can be regarded as holotype for . phyteumatis-orbiculare (Art. 9.1, Note 1).
(6) Uredo bifrons DC., in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 229, 1805.
Holotype: France, on Rumex crispus [patience crépue], ex herb. de Candolle (G00242484).
= Uredo rumicis Schumach., Enum. pl. 2: 231, 1803.
 Uromyces rumicis (Schumach.) G. Winter, Renabh. Krypt.-Fl., Edn 2, 1.1: 145, [1884] 1881.

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Notes: The cited collection is the only specimen deposited in de Candolle’s herbarium that is in
agreement with the protologue. Therefore, it can be regarded as holotype (Art. 9.1, Note 1). The type
material has been examined and turned out to be Uromyces rumicis.
(7) Uredo caprearum DC., [as ‘capræarum’], Fl. franç., Edn 3, 5/6: 80, 1815.
Lectotype (designated here, MycoBank, MBT10006427): “Uredo farinosa”, on Salix caprea, 1806,
without locality, date and collector (G00262487).
 Caeoma caprearum (DC.) Schltdl., Fl. berol. 2: 124, 1824.
 Erysibe caprearum (DC.) Wallr., Fl. crypt. Germ. 2: 204, 1833.
 Podosporium caprearum (DC.) Lév. [as “capraearum”], Ann. Sci. Nat., Bot., Sér. 3, 8: 374, 1847.
 Podocystis caprearum (DC.) Fr. [as “capraearum”], Summa Veg. Scand., Sectio Post.: 512, 1849.
 Lecythea caprearum (DC.) Berk., Outl. Brit. Fung.: 334, 1860.
 Melampsora caprearum (DC.) Thüm., Mitt. Forstl. Versuchswesen Oesterr. 2(1): 34, 1879.
= Uredo farinosa Pers. [. salicis-capreae], Syn. meth. fung. 1: 217, 1801, nom. sanct. Lectotype
(designated here, MycoBank, MBT10006428): Uredo on Salix caprea, without locality, date and
collector, herb. Persoon, no. 91026447 (L0114354).
 Lecythea farinosa (Pers.) Lév., Ann. Sci. Nat., Bot., Sér. 3, 8: 374, 1847, nom. inval. (Art. 35.2).
 Melampsora farinosa (Pers.) J. Schröt., in Cohn, Krypt.-Fl. Schlesien 3.1(17–24): 360, 1887.
 Melampsora salicis-capreae (Pers.) G. Winter, Rabenh. Krypt.-Fl., Edn 2, 1.1: 239, [1884] 1881.
= Sclerotium salicinum Pers., in Mougeot & Nestler, Sirp. Crypt. Vog.-Rhen., Fasc. IV, no. 386, 1813,
nom. nud.
 Sclerotium salicinum Pers. ex DC., Fl. franç., Edn 3, 5/6: 114, 1815. [Syntype: France, on Salix
caprea, Mougeot & Nestler, Sirp. Crypt. Vog.-Rhen. 386 (F0170751F).]
 Xyloma salicinum (Pers. ex DC.) Duby, Bot. Gall. 2: 875, 1830, nom. illeg. (Art. 53.1), non Pers.,
1794.
 Melampsora salicina (Pers. ex DC.) Lév., Ann. Sci. Nat., Bot., Ser. 3, 8: 375, 1847, nom. inval.
(Art. 35.5).
 Melampsora salicina (Pers. ex DC.) Tul., Ann. Sci. Nat., Bot., Ser. 4, 2: 98, 1854.
 Melampsora salicina (Pers. ex DC.) Rabenh., Klotzschii Herb. Viv. Mycol., Ed. Nov., Cent. V: no.
494, 1857, isonym (Art. 6, Note 2).
= Xyloma frustulatum Fr., Observ. mycol. 2: 358, 1818.
= Epitea fenestrata Bonord., Abh. Naturf. Ges. Halle 5: 203, 1860 [also, in Rabenh., Fungi Eur. Exs.
(Klotzschii Herb. Viv. Mycol. Continuatio, Ed. Nova, Ser. Sec.), Cent. 2: no. 189, 1860; Bot.
Zeitung 18: 175, 1860; Flora 43: 749, 1860]. Lectotype (designated here, MycoBank,
MBT10006429): Germany, Guestphalia, on Salix caprea, Bonorden, Rabenh., Fungi Eur. Exs. 189
(HAL, s.n.).
= Entyloma salicis P. Karst., Meddn Soc. Fauna Flora Fenn. 14: 103, 1887 [type host: Salix caprea].
= Melampsora laricis-caprearum Kleb., Forstl.-Naturw. Zeitschr. 6: 469, 1897.
Notes: The nomenclature of this species is complex and in need of some explanations. (1) Melampsora
farinosa, with the sanctioned name Uredo farinosa as basionym, is the valid name for the species,
which is usually referred to as Melampsora caprearum and M. laricis-caprearum, respectively. Jørstad
(1958: 7) examined the original collections of U. farinosa in Persoon’s herbarium. “No. 91026447” is
a collection on Salix caprea, which contains uredinia and telia. Jørstad (l.c.) confirmed its identity as
Melampsora caprearum (M. laricis-caprearum). This specimen is designated as lectotype.
(2) The designated lectotype material of Uredo caprearum from de Candolle’s herbarium has been re-
examined and confirmed as M. farinosa (uredinia hypophyllous, 1–2.5 mm diam., orange, paraphyses
numerous, capitate, capitulae 15–25 µm diam., wall up to 5 µm thick at the apex, urediniospores 13–
22 µm diam., loosely verruculose, distance between verrucae 1–3 µm). The name Melampsora
caprearum (DC.) Thüm. (Thümen 1879), based on Uredo caprearum DC., is sometimes cited as new
species name ascribed to de Thümen. Thümen (l.c.) cited this name as “Melampsora caprearum
Thüm. nov. sp.”, i.e., this name could be interpreted as new name introduce for the teleo- and
holomorph, respectively (according to Art. F.8.1). He cited several synonyms for the uredinial stage
(“stylospores”), including Uredo farinosa and U. caprearum, as synonyms. However, he also cited the
valid name Sclerotium salicinum Pers. ex DC. and several other names as synonyms for the
teleomorph, which would render the name Melampsora caprearum illegitimate (Art. 52.1). Therefore,
this name can only be considered a valid new combination, based on Uredo caprearum, as done in

57
Index fungorum, which is possible since all requirements for a valid publication of a new combination
are given. Thümen (1879a) cited other names in this publication as “nov. sp.” as well, such as
Melampsora vitellinae, although he did not describe the teleomorph, indicating a misuse of “nov. sp.”
in this work, which is in favour of interpreting such names as combinations. The original description
of M. caprearum (without synonyms) was later copied and published in “Hedwigia 18: 77, 1879” with
reference to Thümen (1879a). Hence, the name M. caprearum published in Hedwigia is an isonym of
the validly published name M. caprearum published by Thümen (1879a).
Braun & Bensch (2019) cited Epitea fenestrata Bonord. (type host = Salix caprea), based on
uredinia, as synonym of “Melampsora ribesii-epitea”, which was probably not correct. M. ribesii-
epitea may also occur on Salix caprea (Klenke & Scholler 2015), but M. farinosa is the most common
Melampsora on this host. The urediniospores of the two species are rather similar, and a
differentiation between them without telia is difficult. Therefore, it was necessary to clarify this name
by epitypification with a genetically proven epitype, which will be made in a later phylogenetic
publication.
(8) Uredo longicapsula DC., in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 233, 1805.
Holotype: “Aecidium pynolae Gmel.?”, on Populus nigra, 6 Oct. 1791, Chaillet, ex herb. de Candolle
(G00262494).
 Lecythea longicapsula (DC.) Lév., Ann. Sci. Nat., Bot., Sér. 3, 8: 374, 1847, nom. inval (Art. 35.2).
 Epitea longicapsula (DC.) Rabenh., Klotzschii Herb. Viv. Mycol., Edn Nov, Ser. Sec., Cent. 4: no.
384, 1857.
= Sclerotium populneum Pers., Observ. mycol. 2: 25, [1799] 1800.
Type: on Populus nigra. [Possible type: J.F. de Chaillet [910.264-607] (L0117545)].
 Melampsora populnea (Pers.) Lév. [as “populina”], Ann. Sci. Nat., Bot., Sér. 3, 8: 375, 1847, nom.
inval. (Art. 35.2).
 Melampsora populnea (Pers.) P. Karst., Bidr. Känn. Finl. Nat. Folk 31: 53, 1878.
[Melampsora tremulae Tul., Ann. Sci. Nat., Bot., Sér. 4, 2: 95, 1854, nom. illeg. (Art. 52.1).]
= Uredo populina Pers. [], Syn. meth. fung. 1: 219, 1801, nom. sanct. Lectotype (designated here,
MycoBank, MBT10006430): on Populus basamifera, herb. Persoon [910.264-112], “prope Parisios”
(L00119108).
 Melampsora populina (Pers.) Tul., Ann. Sci. Nat., Bot., Ser. 4, 2: 95, 1854.
[Uredo cylindrica F. Strauss, Ann. Wetter. Gesellsch. Ges. Naturk. 2: 92, 1811, nom. illeg. (Art.
52.1).]
[Caeoma cylindricum Schltdl., Fl. berol. 2: 119, 1824, nom. illeg. (Art. 52.1).]
[Caeoma cylindricum (F. Strauss) Link, in Willdenow, Sp. pl., Edn 4, 6(2): 39, 1825, nom. illeg. (Art.
53.1), non Caeoma cylindricum Schltdl., 1824.]
= Sphaeria populi Sowerby, Col. fig. Engl. Fung. Mushr. 3(no. 25): tab. 374, fig. 2, 1802, nom. inval.
(Art. 38.1a).
 Peripherostoma populi Gray [as “(Sowerby) Gray”], Nat. Arr. Brit. Pl. 1: 515, 1821. Lectotype
(designated here, MycoBank, MBT10006431): Sowerby, Col. fig. Engl. Fung. Mushr. 3(no. 25): tab.
374, fig. 2, 1802.
 Phoma populi (Sowerby) Fr., Syst. mycol. 2(2): 547, 1823.
 Melampsora populi (Gray) M. Morelet [as”(Sowerby) M. Morelet”], Cryptog. Mycol. 6(2): 107,
1985, nom. illeg. (Art. 53.1), non M. populi Mont., in Castagne 1851.
= Uredo ovata F. Strauss, Ann. Wetter. Gesellsch. Ges. Naturk. 2: 92, 1811. Neotype (designated
here, MycoBank, MBT10006432): Germany, Baden-Württemberg, Karlsruhe, Daxlanden,
Rappenwört, Bad, on Populus nigra, 2009, M. Scholler (KR-M-0023902).
= Uredo populi H. Mart., Prodr. Fl. Mosq., Edn 2: 231, 1817. Neotype (designated here, Mycobank,
MBT10006433): Russian Empire, Chernigovskaya Guberniya (Chernigov Governorate), Ostra [now
Ukraine, Chernivtsi Oblast, Kitsman Raion, Ostra], on Populus nigra, as “Melampsora laricis-
populina”, 25 Jul. 1908, G. Newodovski, Griby Ross. Herb. 37 (LE 56347).
 Caeoma populi (H. Mart.) Schltdl., Fl. berol. 2: 123, 1824.
 Erysibe populi (H. Mart.) Wallr., Fl. crypt. Germ. 2: 203, 1833.
= Xyloma populinum Duby, Bot. gall. 2: 875, 1830, nom. illeg. (Art. 53.1), non X. populinum Pers,
1801.
 Melampsora populi Mont., Suppl. Cat. Pl. Mars.: 80, 1851.

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= Epitea oblonga Bonord., Abh. Naturf. Ges. Halle 5: 204, 1860 [also, in Rabenh., Fungi Eur. Exs.
(Klotzschii Herb. Viv. Mycol. Continuatio, Ed. Nova, Ser. Sec.), Cent. 2: no. 190, 1860; Bot.
Zeitung 18: 175, 1860; Flora 43: 749, 1860]. Lectotype (designated here, MycoBank,
MBT10006435): Germany, Guestphalia, on Populus sp., Bonorden, Rabenh., Fungi Eur. Exs. 190
(HAL, s.n.).
= Uredo laricis Westend., Bull. Acad. R. Sci. Belg., Cl. Sci., Sér. 2, 11(6): 650, 1861.
 Caeoma laricis (Westend.) R. Hartig, Wichtige Krankheiten der Waldbäume. Beiträge zur
Mykologie und Phytopatologie für Botaniker und Forstmänner: 93, 1874.
 Melampsora laricis (Westend.) R. Hartig, Bot. Centralbl. 23: 362, 1885, and Allg. Forst-
Jagdzeitung 61: 326, 1885.
 Melampsora tremulae f. laricis (Westend.) R. Hartig, Lehrb. Baumkrankh.: 143, 1889.
= Melampsora balsamiferae Thüm., Mycoth. Univ., Cent. 19: no. 1832, 1881. Lectotype (designated
here, MycoBank, MBT10006436): Germany, Bavaria, Bayreuth, May 1876, on Populus
balsamifera, Thüm., Mycoth. Univ. 1832 (HAL, s.n.). Isolectotypes: Thüm., Mycoth. Univ. 1832
(e.g., B, ILL77500, NEB40826, PUR044184.
= Melampsora laricis-populina Kleb., Z. Pflanzenkrankh. 12: 25 (and 43), 1902.
Notes: Type material of Uredo longicapsula has been re-examined and turned out to be Melampsora
populina, as already pointed out by Tulasne (1854). The urediniospores are broad ellipsoid-ovoid to
oblong, 30–45 × 14–20 µm, wall often thickened in the middle, distance between verrucae 1.5–3 µm,
bold towards the apex, paraphyses clavate-capitate, capitulae 15–25 µm diam. There is a single
collection under the name U. longicapsula in de Candolle’s herbarium that corresponds to the
protologue, including a reference to “Aecidium pynolae Gmel.?” This collection can be regarded as
holotype (see Art. 9.1, Note 1). Tulasne (1854) also introduced the new (superfluous) name
Melampsora tremulae (Sclerotium populneum Pers. was cited as synonym). Tulasne (l.c.) cited S.
populneum Pers. (Persoon 1801: 125), in which Populus tremula was cited as host. However, Persoon
(1801) cited Persoon (1800) as original reference in which S. populneum was described on Populus
nigra. In addition, Tulasne (l.c.) referred to Uredo ovata F. Strauss in connection with the uredinial
stage of M. tremulae, which is a species described on Populus nigra and P. tremula.
The nomenclature of the rust fungus commonly referred to as Melampsora laricis-populina is
complicated and has been notoriously confused since the 19th century. Numerous old names are
involved. The nomenclature of this species can only be disentangled on the basis of clarifications of
the typifications of the names involved. The sanctioned name Uredo populina is the oldest valid name
with priority for this rust species. This name must be ascribed to Persoon [Persoon (1801) cited
“Lycoperdon populinum Ehrh., Pl. Crypt. Lin., Dec. 22, no. 220, nom. nud., and Jacquin (1787 [1786]:
t. 9, f. 2, 3). However, Pl. 9 in the latter work exhibits a higher plant, but the name L. populinum is not
listed in this work and it is also not included in the index]. Léveillé (1847: 375) cited the name
“Sclerotium populinum” (probably in error for Sclerotium populneum) under Melampsora, but he
failed to introduce the combination “M. populneum” (or “M. populinum”). The combination M.
populinum was later validly introduced by Tulasne (1854: 95). He cited “Melampsora populinum
Lév.” and listed Uredo populina Pers. as synonym, which fulfils the conditions for a valid
combination. Type collections of Uredo populina preserved in Persoon’s herbarium were examined
and listed by Jørstad (1958: 8). Collection “910.264-112” (on Populus balsamifera), containing
uredinia and telia, and was identified as M. laricis-populina. This collection is designated as lectotype.
Melampsora populi (Gray) M. Morelet is a homonym (nom. illeg., Art. 53.1) of M. populi
Montagne (in Castagne 1851), a name validly published with reference to Xyloma populinum Duby
(1830: 875), non Pers. 1801. The latter name included a reference to Sclerotium populneum (as
“populinum”), described on Populus nigra, which is a synonym of Melampsora populina.
Sphaeria populi Sowerby is an invalid name (nom. nud., Art. 38.1). A description or diagnosis is
lacking and the original illustration is not accompanied by an analysis (Art. 38.7). This name was later
validated as Peripherostoma populi Gray [as “(Sowerby) Gray”, with description]. However, the exact
name of the host species was not given in Sowerby’s original publication, and it cannot be properly
identified on the basis of the original illustration. Gray (1821) did also only cite “Populus” as host.
Therefore, the common interpretation of Melampsora populi as synonym of Melampsora laricis-
populina, as done, for instance, in Index fungorum, required the clarification of the typification of
Peripherostoma populi. This name is lectotypified by the original illustration. However, this lectotype
requires an epitype, including ex-epitype sequence that fixes the application of this name as currently
done, i.e., as synonym of M. populina. This will be done in a later phylogenetic publication.

59
Strauss (1811) introduced the name Uredo ovata for rusts on Populus nigra and P. tremula. The
identity of this name can also only be clarified and determined by typification. The herbarium of F.
Strauss is preserved at herbarium M, but according to a recent information of D. Triebel, curator of the
mycological herbarium in M, authentic material for this name is not preserved. Therefore, we propose
a neotype, viz., a collection on Populus nigra, the first mentioned host, and follow the assumption in
Sydow & Sydow (1915) that U. ovata belongs to M. laricis-populina (now M. populina).
The synonymy of Epitea oblonga was discussed in Braun & Bensch (2019: 13). The lectotype
material of Melampsora balsamiferae perfectly coincides with M. populina (uredinia hypophyllus,
telia epiphyllous, paraphyses to 70 µm long, clavate to capitate, terminal capitulate 12–24 µm diam.,
wall at the apex 3–10 µm thick, urediniospores 30–38 × 15–20(–23) µm, wall 2–3 µm wide,
sometimes wider in the middle, verruculose, distance between verrucae 1.5–3 µm, less verruculose at
the apex or bold or almost so).
The identity of Uredo populi H. Mart., described from Russia on Populus nigra and P. tremula, can
only be clarified by neotypification as well. The description is very meagre, and two Melampsora
species were undoubtedly involved in the original description. We prefer to use Russian neotype
material on Populus nigra, the host cited first in the protologue, belonging to M. populina, the most
common species on this host.
The type material of Uredo laricis is not preserved at BR in herb. Westendorp. There is a single
specimen deposited under the name Caeoma laricis ( Uredo laricis) [Belgium, on Larix decidua,
s.d., G.-D. Westendorp, s.n. (BR5020105858318)], which cannot be considered the holotype.
Melampsora laricis and M. tremulae f. laricis were introduced as combinations based on Uredo laricis
[by reference to Caeoma laricis] (Hartig, in Anonymous 1885, Hartig 1889), and applied to a
Melampsora host-alternating with Populus tremula. However, Hartig in Anonymous (1885) only
obtained uredinia, i.e., telia were not observed and not described, so that the name Melampsora laricis
cannot be ascribed to Hartig as new teleomorph-typified species name, according to Art. F.8.1. The
holotype of Uredo laricis is not preserved and caeomata on Larix decidua are associated with several
Melampsora species, and the morphological differentiation between them is not possible with
certainty. Therefore, the clarification of the identity of Uredo laricis and its homotypic synonyms
requires the neotypification with genetically proven material. I prefer to follow Hartig’s (l.c.)
interpretation that this name is ascribable to M. tremulae (now M. populina). However, to fix this
interpretation, it is necessary to neotypify Uredo laricis with phylogenetically proven (sequenced)
material. This will be made in a later phylogenetic publication.
Klebahn (1902: 43) reported results of his inoculation experiments, but the morphology of
Melampsora laricis-populina was not described. However, the name M. laricis-populina was first
introduced on page 25, where Klebahn discussed differences to M. allii-populina, which constituted a
diagnosis and valid publication of the name M. laricis-populina.
(9) Uredo petasitis DC., in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 236, 1805.
Lectotype (designated here, MycoBank, MBT10006437): France, “fl. fr. 635”, on Petasites hybridus
( Tussilago petasites), ex herb. de Candolle (G00262465).
 Coleosporium petasitis (DC.) Lév., in d’Orbigny, Dict. Univ. Hist. Nat. 12: 786, [1848] 1849, nom.
inval. (Art. 35.5).
 Coleosporium petasitis (DC.) Berk. [as “Lév.”], Outline of British fungology: 333, 1860.
= Peridermium boudieri E. Fisch., Bull. Soc. Bot. Fr. 41: CLXXI, 1895, nom. nud. (Art. 38.1 a).
= Peridermium dietelii G.H. Wagner [as “dieteli”], Z. Pflanzenkrankh. 6: 10, 1896.
Notes: Berkeley (1860) cited “Coleosporium petasitis Lév.” which is a reference to Léveillé, in
d’Orbigny (1849: 786). Léveillé (l.c.) cited Uredo petasitis DC. under Coleosporium, but did not
introduce the combination C. petasitis (Art. 35.5). Therefore, Berkeley’s (1860) citation can be
considered a new combination based on Uredo petasitis, validated by indirect reference (Art. 38.14,
41.3) to de Candolle’s name.
(10) Uredo pinguis DC., in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 235, 1805.
Lectotype (designated here, MycoBank, MBT10006438): France, “fl. fr. 631”, on Rosa gallica (= R.
austriaca), ex herb. de Candolle (G00566472).
= Puccinia mucronata Pers., Neues Mag. Bot. 1: 118, 1794, nom. sanct. (Persoon 1801: 230).
 Phragmidium mucronatum (Pers.) Schltdl., Fl. berol. 2: 156, 1824.

60
Notes: The name Uredo pinguis is in need of lectotypification. It was introduced with reference to two
different hosts, viz., α. Rosae austriacae and . Rosae alpinae. De Candolle’s original material on
Rosa austriaca, deposited at G, is designated as lectotype.
(11) Uredo punctata DC. [α. euphorbiae-helioscopiae], in Lamarck & de Candolle, Fl. franç., Edn 3,
2: 236, 1805.
Lectotype (designated here, MycoBank, MBT10006439): France, “Aecidium; Lycoperdon epiphyllum
Lin.”, on Euphorbia helioscopia, ex herb. de Candolle (G00262467).
 Diplodia punctata (DC.) Lév., in Orbigny, Dict. Univ. Hist. Nat. 12: 779, 1849.
= Uredo helioscopiae Pers., Neues Mag. Bot. 1: 93, 1794.
 Caeoma helioscopiae (Pers.) Schltdl. [as “(DC.) Schltdl.”], Fl. berol. 2: 125, 1824.
 Erysibe helioscopiae (Pers.) Wallr., Fl. crypt. Germ. 2: 205, 1833.
 Melampsora helioscopiae (Pers.) G. Winter, Rabenh. Krypt.-Fl., Edn 2, 1.1: 240, [1884] 1881.
 Uredo euphorbiae-helioscopiae Pers., Syn. meth. fung. 1: 215, 1801, nom. sanct. Neotype
(designated here, MycoBank, MBT10006440): “Uredo euphorbiae var.”, on Euphorbia helioscopia,
J. J. A. Mougeot, ex herb. Persoon 910.264-560 (L 0114215).
 Melampsora euphorbiae-helioscopiae (Pers.) Nannf., in Lundell & Nannf., Fungi Exsiccati Suecici:
no. 1211, 1943.
[Uredo euphorbiae Rebent., Prodr. fl. neomarch.: 354, 1804, nom. illeg. (Art. 52.1)]
[Lecythea euphorbiae (Rebent.) Lév., Ann. Sci. Nat., Bot., Sér. 3, 8: 374, 1847, nom. inval. (Art.
35.2).]
Notes: The status of the name Uredo punctata requires clarification by lectotypification. De Candolle
(1805) introduced this name with three different hosts (α. Euphorbiae-helioscopiae, . Euphorbiae-
pusillae and . Euphorbiae-pelidis). A collection of α. Euphorbiae-helioscopiae on Euphorbia
helioscopia from de Candolle’s herbarium is designated as lectotype.
The taxonomy and circumscription of Melampsora spp. on Euphorbia spp. have been variously
treated. The Melampsora on Euphorbia helioscopia is biologically confined to this host, i.e., in
inoculation experiments, it could not be transferred to Euphorbia peplus and other Euphorbia species
that pertain to M. euphorbiae (Ficinus & C. Schub.) Castagne (Sydow & Sydow 1915), and this rust is
also phylogenetically different from M. euphorbiae (see Maier et al. 2003), which has been taken into
account in the present synonymy of M. euphorbiae-helioscopiae.
According to the current Code (Art. F.3.7), Melampsora euphorbiae-helioscopiae is the correct name
for this fungus, since Uredo euphorbiae-helioscopiae being a sanctioned name, treated as if conserved
against earlier homonyms and competing synonyms [Persoon (1801) cited “Disp. meth. fung. p. 13”
(Persoon 1797: 13) in the protologue, which is a reference to the name Uredo helioscopiae in the latter
work, based on the original publication of this name in Persoon (1794), so that U. helioscopiae has to
be considered a homotypic synonym of U. euphorbiae-helioscopiae]. Nannfeldt’s combination in
Fungi Exs. Suec. was effectively published (see Art. 30.8, Ex. 12). Jørstad (1958: 6) examined
specimens deposited in Persoon’s herbarium under Uredo euphorbiae/helioscopiae/euphorbiae-
helioscipiae and emphasized that suitable type material is not preserved. Therefore, a neotype is
designated, based on a collection examined by Jørstad (l.c.) and confirmed as “Melampsora
euphorbiae (syn. M. helioscopiae)”. The neotype only contains uredinia.
De Candolle, in Lamarck & de Candolle (1805), cited Uredo helioscopiae with reference to Persoon
(1801), who introduced Uredo euphorbiae-helioscopiae with reference to Uredo helioscopiae in
Persoon (1798). Hence, de Candolle (l.c.) indirectly referred to U. helioscopiae Pers. and did not
intend to introduced a new species with the same name. Hence, all combinations based on “U.
helioscopiae DC.”, such as Caeoma helioscopiae refer to U. helioscopiae Pers. as basionym. Uredo
euphorbiae Rebent. is an illegitimate (superfluous) name, since the original description encompassed
Uredo euphorbiae-helioscopiae [a reference to Persoon (1801: 215) was given].
(11) Uredo salicis DC., in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 230, 1805.
[Type: on Salix sp. (“Uredo du saule”), C. Berger, without any further details.] Lectotype (designated
here, Mycobank, MBT10006441): France, on Salix triandra, without locality, date and collector, ex
herb. de Candolle (G00262500).
= Hypodermium (subgen. Uredo) mixtum Link [as “Ur. mixta”], Mag. Gesell. Naturf. Freunde, Berlin
7: 28, 1816 [type host: Salix triandra].

61
 Caeoma mixtum (Link) Schltdl., Fl. berol. 2: 124, 1824.
 Uredo mixta (Link) Duby, Bot. Gall. 2: 895, 1830.
 Erysibe mixta (Link) Wallr., Fl. crypt. Germ. 2: 204, 1833.
 Lecythea mixta (Link) Lév., Ann. Sci. Nat., Bot., Sér. 3, 8: 374, 1847, nom. inval. (Art. 35.2).
 Epitea mixta (Link) Fr., Summa veg. Scand., Sectio Post.: 512, 1849.
 Melampsora mixta (Link) Thüm., Mitt. Forstl. Versuchswesen Oesterr. 2(1): 42, 1879.
= Melampsora amygdalinae Kleb., in Pringsheim, Jahrb. Wiss. Bot. 34: 352, 1900 [type host: Salix
triandra].
Notes: The identity of Uredo salicis remained quite unclear to this day. The identity of this name can
only be pointed out in connection with a clarification of its typification. In the original description, de
Candolle (in de Lamarck & Candolle 1805) cited a single specimen on Salix, collected by C. Berger.
However, de Candolle (l.c.) mentioned C. Berger and names of other collectors or senders in the
protologues of several other names of rust fungi, but the names concerned are often not to be found on
the labels of authentic collections in de Candolle’s herbarium that were recently re-examined. The
collection on Salix triandra, deposited in de Candolle’s herbarium under the name Uredo salicis is,
although without any other data on the label, the only specimen that can be taken into consideration as
original material. It is unclear if de Candolle obtained a single specimen that he used for the
description of U. salicis. Therefore, I do not consider this sample, according to Art. 9.1, Note 1, as
holotype, but prefer to designate it as lectotype. The collection on Salix triandra from de Candolle’s
herbarium, which is designated as lectotype of Uredo salicis, has been examined and identified as
Melampsora, so far usually referred to as M. amygdalinae [uredinial paraphyses capitate, capitate apex
12–22 µm wide, uredospores oblong ellipsoid-ovoid, 18–32 × 12–18 µm, with loosely arranged warts,
distance between warts 1.5–3 µm, apically with fewer or without any verrucae (“bold”)].
Hypodermium (subgen. Uredo) mixtum is another synonym. Link (1816) introduced this name and
cited Uredo salicis as synonym with question mark. Schlechtendal (1824) introduced the new
combination Caeoma mixtum and cited Uredo salicis as genuine synonym (without question mark).
Last but not least, Thümen (1879a) introduced the combination Melampsora mixta and classified Salix
triandra as principal host [this name was introduced as “Melampsora mixta nov. spec.” but it fulfils
the necessary conditions for a new combination; the basionym was cited as synonym and the
teleomorph was not described, so that Art. F.8.1 cannot be applied].
The name Uredo salicis takes priority over Hypodermium (subgen. Uredo) mixtum and Melampsora
amygdalinae. To maintain the established name Melampsora amygdalinae, a proposal to preserve this
name is needed.
(12) Uredo vitellinae DC., in Lamarck & de Candolle, Fl. franç., Edn 3, 2: 231, 1805.
Lectotype (designated here, MycoBank, MBT10006442): on Salix alba var. vitellina ( S. vitellina),
Girod-Chantrans (1802: plate 22, figs. 55, 55’).
 Erysibe vitellinae (DC.) Wallr., Fl. crypt. Germ. 2: 204, 1833.
 Epithea vitellinae (DC.) Fr. Summa veg. Scand.: 512, 1849.
 Melampsora vitellinae (DC.) Thüm., Mitt. Forstl. Versuchswesen Oesterr. 2(1): 43, 1879.
 Melampsora vitellinae (DC.) Thüm. [as “Thüm.”], Hedwigia 18(5): 79, 1879, isonym. (Art. 6, Note
2).
[Caeoma saliceti Schltdl., Fl. berol. 2: 124, 1824, nom. illeg. (Art. 52.1).
 Uredo saliceti (Schltdl.) J. Becker, Fl. Frankfurt, Zweite Abth.: 252, 1828.
 Lecythea saliceti (Schltdl.) Berk., Outl. Brit. Fung.: 334, 1860.]
= Uredo orbicularis Mart., Fl. crypt. erlang.: 318, 1817 [type host: Salix alba].
= Melampsora salicis-albae Kleb., Jahrb. Wissen. Bot. 35: 679, 1901, nom. inval. (Art. 36.1).
= Melampsora allii-salicis-albae Kleb. [as “nom. nov.”], Z. Pflanzenkrankh. 12: 19, 1902.
Notes: De Candolle (in Lamarck & de Candolle1805) introduced the name Uredo vitellinae for an
uredinial stage on “saule osier” (Salix vitellina  S. alba var. vitellina). De Candolle (l.c.) based this
name on an illustration published by Girod-Chantrans (1802), who described and discussed a rust
(rouille) on Salix vitellina (Girod-Chantrans 1802: 131). Girod-Chantrans (1802: Pl. 22, fig. 55) might
refer to a leaf of Salix alba var. vitellina, and Fig. 55’ can be interpreted as uredinia and
urediniospores of a Melampsora. They are not globose, but rather ellipsoid-oblong, which is in
agreement with those of Melampsora on Salix alba, usually referred to as M. salicis-albae, which is,
however, an invalid name, published as “ad int.” (Art. 36.1). Additional specimens were not cited, so

62
that only the original illustration can be used for lectotypification. However, the identity of U.
vitellinae can only be pointed out by clarification of its typification. The lectotype (Girod-Chantrans
1802: Pl. 22, fig. 55) is not sufficient for a proper identification of this name. Therefore, the lectotype
must be supplemented by an epitype. This will later be accomplished in another publication with
corresponding, sequenced material.
In de Candolle’s herbarium, there several specimens, collected after 1805, which are deposited under
Uredo vitellinae and a single undated uredinial sample (without locality and collector) on Salix
viminalis (G00261342), which is morphologically identical to Melampsora epitea Thüm. (uredinia
about 0.5 mm diam. or somewhat larger, paraphyses capitate, capitulae 20–30 µm diam.,
urediniospores globose, 16–24 µm diam., regularly verrucose, distance between verrucae 1.5–3 µm).
Thümen (1879a) introduced the new combination Melampsora vitellinae, described “stylospores”
(urediniospores), and assigned collections on Salix fragilis, S. lucida, S. pentandra, and S. vitellina to
this name, but emphasized that S. vitellina being the main host. Telia and teliospores were not
described. Therefore, the name M. vitellinae, although described as “nov. spec.”, can only be
considered a new combination based on Uredo vitellinae, which was cited as synonym, as also
interpreted in Index fungorum and MycoBank. In Thümen (1879b), all descriptions of species from
Thümen (1879a) were repeated with reference to the latter publication, but without citing any
synonyms, which constitutes an indirect reference to Uredo vitellinae (Art. 43.1). Sydow & Sydow
(1915) cited Uredo vitellinae as synonym of Melampsora laricis-pentandrae, but only with question
mark.
Caeoma saliceti (Schlechtendal 1824) is an illegitimate (superfluous) name, according to Art. 52.1,
since Uredo vitellinae was cited as synonym.
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Address of the author
Uwe Braun, Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und Botanischer Garten,
Neuwerk 21, 06099 Halle (Saale), Germany.
(E-mail: uwe.braun@botanik.uni-halle.de)