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THE BRITISH SPECIES OF PUCCINIA INCLUDEDUNDER "P. SYNGENESIARUM" WITH NOTES UPON

THE BRITISH RUST FUNGI OCCURRINGON THISTLES

By MALCOLM WILSONDepartment ofMycology, Unioersity ofEdinburgh

THERE has been considerable doubt regarding the identity of therusts known by the older British mycologists as Puccinia Syngenesiarum.*Two authorities have been given for this species, Link and Corda.

(I) P. Syngenesarum Link in Linn. Sp. Plant. ed. IV, 6, (2), p. 74(1825).

P. conglomerata Schmidt & Kunze, exsicc. n. 191.Uredo conglomerata Strauss, Welt. Ann. 2. p. 100.

This is stated by Link to occur on T ussilago alpina Kunze and onCentaurea alpine Ehrenb. inGermany. These two hosts have beenproved to be identical and are now called Homogynes alpina. The rustis known as P. conglomerata (Strauss) Kze. & Schm. It is a well­recognized species possessing teleutospores only, which are smooth,generally attenuated at base and apex, with the apex usually un­thickened and with very short hyaline pedicels. A description withreferences to literature, iconography, synonymy and list of exsiccatais given by Sydow (1902, p. 99). P. Senecionis Lib. and P. expanseLink were at one time included in P. conglomerata but are now recog­nized as distinct species. There is no evidence that P. conglomerate as atpresent defined is a British species.

(2) P. Syngenesarum Corda, leones IV, p. 16, Taf. rv, fig. 53 (1840).Corda adds "Link. Spec. II, p. 74, part", and states that the speciesoccurs on Cirsium lanceolatum, Onopordon Acanthium and many otherSynantherae.

He adds that the large, elongated, almost pulvinate and pulveru­lent sori reach often nearly a line in length and up to one-quarter ofa line in width and that the surrounding leaf surface with its hairbecomes powdered with the spores. The spores possess a very short,thin, colourless stalk which is often pointed. Corda gives two figures;oneshowstheleaf, apparently at natural size, with elongated sori about

* The specific epithet has been spelt in several ways: S;ngenesiarum, S;'ngenesarumand S;ngenesearum. The first method, from S;ngenesia L., is regarded here as correctand is used in this note. In referring to special papers however or books the spellingused there is retained.

Rust Fungi occurring on Thistles. M. Wilson 2452-3 mm . long and I mm. wide; the second shows the teleutosporeswith wall of equal thickness all round, no papilla, very slight con­striction, rounded above and below, colourless pedicel more or lesspointed at the base and sometimes attached obliquely.

It is noteworthy that both P. Syngenesarum Link and P. Syngene­sarum Corda agree in possessing smooth, slightly constricted teleuto­spores with the wall of equal thickness all round and very shorthyaline pedicels.

The descriptions of P. Syngenesiarum given by Johnston (1831,P: 197), Berkeley (1836, p. 365) and Cooke (1871, p. 499; r878, p. 206)are generally similar to the accounts given by Link and Corda.Johnston gives Link as the authority and states that the species occurson the leaves of thistles and of the goat's-beard, Berkeley also givesLink as authority; he states that he has not seen the fungus onthistles and refers to Johnston. Cooke gives Link as the authority butalso refers to Corda; he quotes P. CirsiiFckl. as a synonym. He statesthat the species is common on thistles in autumn (1871, p. 499; 1878,p. 206). Plowright (1889) gives P. Syngenesiarum Link as a synonymfor P. Tragopogi and quotes "Johnst. Flor. Berw. II, p. 197, in part".

Up to this stage all the authors agree in their publications thatP. Syngenesiaruni possesses tcleutospores with very short pediccls. It~ay be assumed that this was the general opinion until the publicationin 1889 of Plowright's British Uredineae and Ustilagineae.

When however the specimens of P. Syngenesiarum collected byJohnston, Vize and Cooke are examined, a different state of affairs isdiscovered. Eighteen specimens from the herbaria at Kew, Edin­burgh, Birmingham (Plowright Herbarium and Grove Collection)and Berwick (Johnston) have been examined; these specimens werecollected between 183I and 1885. The oldest specimen examined,*that from Johnston's herbarium at Berwick, is found to be P. Cardu­orum Jacky, on Carduus ctispus L. As P. Carduorum possesses small soriand the tcleutospores have short hyaline pedicels it docs generallyagree with the description given for P. Syngenesianim. Two of .thcspecimens were collected by Vize; one is P. Carduorum on Carduusnutans L., and the second P. Le Monnieriana Maire on Cnicus palustrisWilld. One specimen collected by Miss Jelly in 1878 is also P. LeMonnieriana on C. palustris. Of the fourteen specimens collected ornamed by Cooke, one is P. Carduorum on Carduus crispus and thirteenare P. Le Monnieriana on C. palustris.

In the majority of the specimens of P. Syngenesiarum, the name ofthe host plant is not given and, since most of the specimens consist of

* There is no date on Johnston's specimen in the herbarium at Berwick but as areference is given on the sheet to his Flora it must have been placed in this collectionsubsequently to 1831. Johnston died in 1855 and thus the date of the specimenmust be between 1831 and 1855.

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246 Transactions British Mycological Society

isolated leaves or portions of leaves, it has been very difficult todetermine the identity of the hosts. The differences between the leavesof Cnicus lanceolalus and C..palustris are, however, well marked. InC. lanceolatus stout spines are found on the margin of the leaf andsimilar spines ofsmaller size occur on the upper surface; a few white,cottony hairs are also found on the upper surface, especially along-th­midrib and veins and similar hairs are abundant on the lower surface.In C. palusttis, spines are found only on the leaf margin; scattered,multicellular hairs, which have a glistening appearance in the driedspecimen, are found scattered over both upp~r and under surfaces,especially near the veins; these somewhat resemble a string ofbeads inappearance, owing to the constrictions between the cells.

The teleutosori of P. Le Monnieriana are fairly large, the teleuto­spores are clavate with long, coloured, persistent pedicels and thusthirteen out of fourteen of Cooke's specimens do not agree with hisdescription of P. Syngenesiarum (1878, p. 206) in which the sori aredescribed as minute, the teleutospores having short pedicels.

It is interesting to note that in the packet containing one of Cooke'sspecimens there is a portion of a letter written by the correspondentwho sent the specimen; the signature has been cut away. This corre­.spondent writes: "What is this enclosed Puccinia on Carduus? It isunlike any of the described species on the Compositae as it has a verylong peduncle." This specimen was, however, labelled P. Syngenesi­arum by Cooke.

Cooke, in his general account of P. Syngenesiarum states (1878,p. 62): "No species in the entire genus makes so prominent an appear­ance as the one on the radical leaves of the spear thistle (Carduuslanceolalus). This latter plant is exceedingly abundant, and so is itsparasite (Puccinia syngenesiarum, Lk.}, From the month ofJuly till thefrosts set in we may be almost certain of finding specimens in anywood. The leaves have a paler roundish spot, from one-twelfth to one­fourth ofan inch in diameter, on the upper surface, and a correspond­ing dark brown raised spot on the under surface, caused by an aggre­gation of pustules, forming a large compound pustule, often partlycovered with the epidermis. The individual pustules are small, butthis aggregate mode of growth gives the clusters great prominence,and therefore they are not easily overlooked (plate IV, fig. 63).Although not confined to this species of thistle, we have not yetfound this Puccinia on any other plant. The spores are elliptical,rather elongated, constricted, and without spines (fig. 64)."

If, in this account, Cnicus palustris is substituted for C. lanceolalus asthe name of the host, a very good description of P. Le Monnieriana isproduced; the observation that the rust occurs on the radical leaves isparticularly interesting, for this position is very characteristic ofP. LeMonnierlana and has been specially noted by Maire (1913); the pale

Rust Fungi occurring on Thistles. M. Wilson 247

roundish spot on the upper surface of the leaf opposite the hypo­phyllous sorus is also distinctive of this species. From a considerationof Cooke's account, together with the facts previously mentioned, itappears very probable that his identification of the host was incorrectand that the plant in question was Cnicus palustris, not C. lanceolatus.Additional evidence for this conclusion is given by an examination ofhis fig. 63, PI. IV, in which a portion of the leaf, bearing sori isshown; the leaf depicted agrees better with C. palustris than with C.lanceolatus and the sori closely resemble those of P. Le Monnieriana.Also it may be noted that C. lanceolalus does not commonly grow inwoods while C. palustris is often found in open places in damp woods.Cooke's fig. 64, PI. IV depicts two teleutospores; these, as they arehardly clavate and have short colourless pedicels and no apical thick­ening, do not agree with those of P. Le Monnieriana but more nearlywith the description of P. Syngenesiarum.

During August 1939 plants ofCnicus palusttiswere. collected in WyreForest, Worcestershire, in which the radical leaves were infected withP. Le Monnieriana and die cauline leaves with P. Cirsii. Cooke mayhave collected similarly infected plants and drawn the sori from aradical leaf, and the teleutospores from a cauline leaf; or his drawingof the teleutospores may have been influenced by his description ofP. Syngenesiarum (1878, p. 206).

Knowing that Cooke's P. Syngenesiarum consisted almost entirely ofP. Le Monnieriana on Cnicus palusttis it is easy to trace the influence ofhis mistakes on subsequent investigators. These latter appear to havebased their work on the assumption that P. Syngengsiarum of Cookeconsisted for the most part of a rust growing on Cnicus lanceolatus inwhich the teleutospores possessed an upper cell with the wall thick­ened at the apex and long, persistent, coloured pedicels.

Plowright (1889, p. 216) described a new species P. Cardui onCnicus lanceolalus and Carduus crispus, of which the description agreesalmost exactly with that of P. Le Monnieriana Maire. This is stated togrow on Cnicus lanceolatus and Carduus crispus. He gave as synonyms:

Puccinia Syngenesiarum, Link. Johnst. Flor. Berw. II, p. 197; Berk.Eng. Flor. v, p. 365; Cooke, Hdbk. p. 499; Micro. Fungi, 4th ed.p. 206, t. IV, figs. 63, 64;

Puccinia Cirsii FckI. Exs. No. 340 (?) ;but stated" It is clearly not the plant described by Link (Sp. Plant.VI, pt. II, p. 74) which has very short pedicels. It may be Fuckel'sP. Cirsii."Plowright thus clearly indicated that although his specieswas not P. Syngenesiarum Link it was the P. Syngenesiarum of Cooke andothers. Up to the present, endeavours to obtain Plowright's typespecimen have failed. Specimens of P. Cardui Plowr, have, however,been examined from Kew and Birmingham. That from Kew islabelled "Herb. C. Crossland, on Carduus sp.? Isle of Wight, per J. F

248 Transactions Britisb Mycological Society

Rayner, Southampton, Aug. 29/07." It is P. Le Monnieriana onCnicus palustris. Two specimens from Birmingham labelled "PucciniaCardui on Cnicus lanceolatus, Randan 'Woods" (one in the handwritingof W. B. Grove), are also P. Le Monnieriana on Cnicus palustris. Itappears, therefore, that both Plowright and Grove followed Cooke intheir incorrect determination of the host plant; possibly they acceptedCooke's statements and specimens without further investigation.

Sydow (1902, p. 58) quoted P. Cardui as a synonym for P. Cnici­oleracei Pers. and stated that this rust is found on Cirsium lanceolatus andCarduus crispus (?) as well as on Cirsium ochroleucum and C. oleraceum,apparently relying on Plowright's statements for the two formerspecies. Fischer (19°4, p. 292) stated that P. Cnici-oleracei occurs onCirsium ochroleucum, C. oleraceum, C. lanceolatum and Carduus crispus"(nach P. und H. Sydow) ", but records it in Switzerland only onCirsium oleraceum.

Grove (1913, p. 144) under his description of P. Cnici-oleracei Pers.states that" It is doubtful if Puccinia Syngenesiarum (Cooke, Handbook,p. 499; Micro. Fung, p. 206) belongs entirely here, as the figure in thelatter work (PI. IV, fig. 64) docs not give the true form of the teleuto­spore; but the majority of the specimens issued by him under thatname are this species." Grove quotes P. Cardui Plowr. as a synonymfor P. Cnici-oleracei Pers., and gives as host plants Cnicus Ianceolatus. and,doubtfully, Carduus crispus. This is not surprising, for there is a greatresemblance. between P. Cnici-oleracei and P. Le Monnieriana and, asalready shown, it is highly probable that P. Cardui Plowr. is identicalwith P. Le Monnieriana.

It may be concluded therefore that Cooke's misstatements, madebetween 1864 and 1885, and his incorrectly determined specimens,strongly influenced investigators during the subsequent thirty-fiveyears and at length led to the erroneous statements made by Grove inconnexion with P. Cnici-oleracei. No British specimens labelled P.Cnici-oleracei have been found in the herbaria at Kew, Edinburgh,Birmingham or Berwick; Ramsbottom (1913) includes this species inhis list of British Uredinales but gives P. Cardui Plowr. as a synonym.

Liro (1908, p. 396) has stated that in Finland, P. Cnici-oleraceioccurs on Cnicus heterophyllus Willd., C. oleraceus and the hybrid C.oleraceus x heterophyllus, and that the rust on these three hosts is identicalwith P. Andersoni B. & Br. Sydow (1904, p. 856) quotes this statementand appears to agree with it. The rust which occurs in this country onCnicus heterophyllus (Plowright, 1889, p. 204; Grove, 1913, p. 146) hasup to the present been regarded as the distinct species P. AndersoniB. & Br., but in view of Liro's discovery it must now be regarded asPuccinia Cnici-oleracei Pers.

Puccinia Cardui-pycnocephali Sydow has now been recorded byHadden (1930) in Somerset, Ellis (1933-4) in Norfolk, Wilson (1934,

Rust Fungi occurring on Thistles. M. Wilson 249

P: 369) in East Lothian and Berwickshire and Mayfield (1935) inSuffolk, in addition to the records given by Grove (1913, p. 142).P. galatica Sydow, at first separated from P. Cardui-pycnocephali, hasbeen recognized as identical with it by Sydow (1904, p. 852).

Puccinia Cirsii-lanceolati Schroet., in Cohn, Krypt.-Fl. Schles. III,

pt. I, p. 317 (1887), should be known as P. Cnici Mart., Fl. Mosq.p. 226 (1817) as pointed out by Arthur (1934-). It has been foundrecently on Cnicus eriophorus Roth., by Ellis and Rhodes near Oxford(Ellis in litt.).

Puccinia Cirsii Lasch has now been found on Cnicus palustris Willd.,on C. pratensis Willd., in Galway (Grove, 1913) and in Surrey, on C.acaulis in Norfolk (Ellis, 1933-4) and in Wiltshire and on C. hetero­phyllus in Scotland (Wilson, 1934).

Puccinia Le Monnieriana Maire is stated by Maire (19 I 3) to beidentical with P. Cirsiorum var. 2 Cirsii-palustris Desm. Pl. Crypt.France, ed. I, no. 557.

Puccinia Cirsiorum var, 1 Cirsii-oleracei is stated by Desmazieres to bePuccinia Cnici-oleracei.

The valid name for P. Le Monnieriana should therefore be P. Cirsii­palustris (Desm.) comb.nov, This species is now known from Middle­sex, Kent, Surrey, Hampshire, Norfolk, Suffolk, Cambridge,Worcestershire and from several localities in Scotland.

Puccinia suaveolens (Pers.) Rostr., common on Cnicus arvensis Hoffm.,has been found on Cnicus arvenis Hoffm., var. setosus Bess., near Inver­ness, by Dr ]. A. Macdonald (in litt.). As pointed out by Magnus(1903) Puccinia suaueolens (Pers.) Rostr, Forh, skand. Natuif. Kopen­hagen, p. 339,1873 is the valid name for this species and not P.obtegens (Link) Tul.

My thanks are due to the following for the loan of specimens andfor facilities for the examination of material: the Director of theRoyal Botanic Gardens, Kew; the Regius Keeper of the RoyalBotanic Garden, Edinburgh; the Professor of Botany, the University,Birmingham; the Municipal Authority, Berwick-upon-Tweed. Iwish to thank Miss E. M. Wakefield, Dr C. G. C. Chesters, andDr G. R. Bisby for help and advice in the preparation of the paper.

SUMMARY

I. Eighteen specimens of Puccinia Syngenesiarum have been exam­ined, and of them fifteen have been found to be P. Le MonnierianaMaire on Cnicus palustris.

2. The mistaken identification by Cooke of C. palusitis for C.lanceolatus as the host plant of Puccinia Syngenesiarum has very probablyled to the erroneous citations of Cnicus lanceolatus as a host plant of

250 Transactions British Mycological SocietyPuccinia Cardui Plowr, by Plowright and as a host of P. Cnici-oleraceiPers. by Grove.

3. Although the type specimen of P. Cardui Plowr. has not beenexamined, this species is almost certainly identical with P. LeMonnieriana Maire.

4. Notes of the nomenclature and distribution of the followingspecies are given: P. Cnici-oleracei Pers. (P. Andersoni B. & Br.), P.Cardui-pycnocephaliSydow, P. Cnici Mart. (P. Cirsii-lanceolati Schroet.),P. Cirsii Lasch, P. Cirsii-palustris (Desm.) comb.nov. (P. Le Monnieri­ana Maire), P. suaveolens (Pers.) Rostr. (P. obtegens (Link) Tul.).

REFERENCES

ARTHUR, J. c. (1934). Manual of the Rusts in UnitedStales and Canada. Lafayette,Indiana, U.S.A. Purdue Research Foundation.

BERKELEY, M.]. (1836). In W.]. Hooker, The English Flora, v, Pt. II. London.COOKE, M. C. (1871). Handbook ofBritish Fungi. London.-- (1878). Microscopic Fungi. 4th ed. London.

ELLIS, E. A. (I933-4). "The Flora of Norfolk. Rust Fungi {Uredinales)." Trans.Naif. Norwich Nat. Soc. XIII, 489.

FISCHER, E. (1904). In Beitrage zur Kryptogamen Flora der Schuieie. rr, Heft. 2. DieUredineen der Schweiz. Bern: K.]. Wyss.

GROVE, W. B. (1913). The British Rust Fungi (Uredinales). Camb. Univ, Press.HADDEN, N. G. (1930), "Uredineae of \Vest Somerset." J. Bot. LVIII, 37-9.] OHNSTON, G. (I 83 I). A Flora ofBerwick-upon-Tweed. Vol. II, Cryptogamous Plants.

Edinburgh: Carfrae and Son.LIRa, ].]. (1908). Uredineae Fennicae. Finlands Roststampar. Helsingfors: Finsku

Litteratur-Sallakapets.l\fAGNUS, P. (1903). "Bemerkungen zur Benennung einiger Uredineen in P. & H.

Sydows' Mon0ll,raphia Uredinearum." Hedwigia, XLII, 305.MAIRE, R. (1913). 'Etudes mycologique, I." Ann. Mycol, XI, 352.MAYFIELD, A. (1935). "The Rust Fungi of Suffolk." Trans-.Sziffolk Nat. Soc. III, 1.PLOWRIGHT, C. B. (1889). A Monograph. of the British Uredineae and Ustilagineae.

London: Kegan Paul, Trench and Co.RAMSBOTTOM,]. (I9I3). "Some notes in the history of the classification of the

Uredinalcs," Trans. Brit: Mycol, Soc. IV, 77-1°5.SYDOW, P. & H. (1902-4). Monographia Uredinearum, Vol. I. Lipsiae: Borntraeger,WILSON, M. (1934). "The Distribution of the Uredineae in Scotland." Trans. Bot.

Soc. Edinb. XXXI, 345-449.

(Acceptedfor publication 8 April 1940)